Monday, August 8, 2022

Vampirism and bloodlust


Ishbosheth is slain (Wikicommons – Maciejowski Bible)


Before the State monopoly on violence, an adult male was expected to spill another man’s blood in the course of life. Such action would be authenticated by the sight, feel, and taste of that blood.



Vampirism is the desire to see, feel, and taste blood. Today, we encounter it in horror movies or Gothic fiction, yet it does exist in real life. A “vampire” derives intense pleasure, bordering on sexual excitement, from the sight, feel, and taste of blood (Jaffé and DiCataldo 1994; Vanden Bergh and Kelly 1964). The following is a classic case:


In 1978, during a two-day rampage in the Mayenne region of France, a 39-year-old man attempted to rape a preadolescent girl, also biting her deeply in the neck, murdered an elderly man whose blood he drank and whose leg he partially devoured, killed a cow by bleeding it to death, murdered a married couple of farmers, and almost succeeded in doing the same with their farm hand. (Jaffé and DiCataldo 1994)


Most of the literature on vampirism comes from societies where the State has long held a monopoly over the use of violence and where non-State violence has long been criminalized and even pathologized. In many parts of the world, however, that monopoly is either recent or ineffective. The average man is still expected to use violence to defend himself and his family against threats that may seem trivial in a State-pacified society.


We need a cross-cultural study of the desire to shed blood. An initial step in that direction was taken by Frantz Fanon, who worked as a hospital psychiatrist in Algeria. He described vampirism as a frequent characteristic of murder cases in that country. The Algerian murderer “needs to feel the heat of blood and steep himself in his victim’s blood. […] A number of magistrates even go so far as to say that killing a man for an Algerian means first and foremost slitting his throat” (Fanon 2004[1963], p. 222).


Additional cross-cultural perspective has been provided by two recent papers. One of them is a case report from Sri Lanka:


A 20-year old single, unemployed male was referred from a drug rehabilitation center to the psychiatry clinic. He presented with poor anger control, impulsive behavior and the urge to drink blood, against a background of multiple substance dependence. He had been adopted in his early childhood and there was no childhood features to suggest developmental delays, hyperactivity, impulsivity, or conduct disorder.


[…] Although he experienced a sense of satisfaction after ingestion of blood, this act was not associated with obsessions, delusions, hallucinations, sexual gratification or paraphilic behaviour. He did not have any other psychiatric illnesses. (Adicaram et al. 2021)


The other paper presents two case reports from Turkey. In that country, vampirism usually takes the form of self-mutilation, if only because personal bloodletting is less likely to invite legal retribution. The authors describe it as following a stereotypical behavioral sequence:


We propose the term "hemomania" to describe an impulse control disorder characterized by impaired functioning due to at least one of the following urges: seeing one's own blood, self-bloodletting, and tasting/drinking one's own blood. We argue that hemomania progresses from an urge to see one's own blood to the urge to drink it (Kandeğer et al. 2021).


The “vampire” responds positively to the sight of blood and is thus driven to spill more blood and ultimately bathe in it and taste it. If this is indeed an impulse-control disorder, it should exist in many apparently normal people, among whom it would be unexpressed and under strong inhibition.


A desire to shed blood may have been much more common before the State monopolized the use of violence—at a time when an adult male was expected to spill another man’s blood in the course of life. In that context, it would be counterproductive to feel nauseated. In fact, one should feel excited. And the final triumph over an adversary would be authenticated by the sight, feel, and taste of that man’s blood.


We still have a word for that: “bloodlust.” There is also the word “bloodthirsty.” Today, we hear and say those words without fully understanding their original meaning. They refer to a mental state that used to be common in another time, but which has since been expunged from normal life … to the point that we now see it as weird and pathological.


A Middle English ballad describes the pleasure that a group of men felt when drinking the blood of a freshly killed deer:


They eat of the flesh, and they drank of the blood,

And the blood it was so sweet,

Which caused Johny and his bloody hounds

To fall in a deep sleep. (Haughey 2011, p. 350)


Those men were breaking a taboo against drinking an animal’s blood or eating its bloody flesh. That taboo went back to Anglo-Saxon times, when meals would bring many men together around the same table. It was feared that consumption of animal blood would excite the male mind and lead to violence, murder and, ultimately, consumption of human blood:


From this savage sharing of raw food with dogs, it is a short logical leap to cannibalism, the ultimate food taboo, for once one is able to devour bloody flesh, one has lost inhibitions concerning food. […] Johny Cock eats raw meat with his dogs, many Robin Hood ballads fixate on the sublimated violence in overblown feast scenes, and uncouth outlaw heroes like Hereward, Gamelyn, and Fulk Fitz Waryn repeatedly break taboos against mixing raw human blood with their meals when they bleed on their plates or tables and insist on continuing their feasts. (Haughey 2011, pp. 29-30)


Many cultural traditions insist on the removal of blood from flesh before it can be eaten. This taboo is described in the Hebrew Scriptures:


But you must not eat meat that has its lifeblood still in it. And for your lifeblood I will surely demand an accounting. I will demand an accounting from every animal. And from each human being, too, I will demand an accounting for the life of another human being. Whoever sheds human blood, by humans shall their blood be shed.

Genesis 9:4-6


With the rise of State societies, male violence became criminalized in most cases, with the notable exceptions of self-defense and war. Those new circumstances favored a different sort of man, one who would react negatively to the sight of blood. With the marginalization of bloodthirsty individuals, and their gradual removal from the gene pool, there was likewise a removal of bloodlust from real life.


Today, bloodlust survives as a deactivated behavior that normally remains dormant. This is the situation that prevails in long-pacified societies: vampirism has literally become pathological—it is reactivated only by environmental or genetic accidents that cause many other pathologies. The “vampire” looks and acts like a freak.


This is less true in societies that have been pacified more recently. The “vampire” seems more normal and shows fewer signs of mental disorder.





Adicaram, D.R.S., Wijayamunige, E.S. and Arambepola, S.C.A., 2021. Vampires! Do they exist? A case of clinical vampirism. Sri Lanka Journal of Psychiatry 12(2): 38-40.


Fanon, F. (2004[1963]). The Wretched of the Earth. New York: Grove Press.


Haughey, S. (2011). The 'Bestli' Outlaw: Wilderness and Exile in Old and Middle English Literature. PhD dissertation, Cornell University.


Jaffé, P. D., and F. DiCataldo. (1994). Clinical vampirism: Blending myth and reality. Bulletin of the American Academy of Psychiatry & the Law 22(4): 533–544.


Kandeğer, A., F. Ekici, and Y. Selvi (2021). From the urge to see one’s own blood to the urge to drink it: Can hemomania be specified as an impulse control disorder? Two case reports. Journal of Addictive Diseases 39(4): 570-574.


Vanden Bergh, R.L.,and J.F. Kelly. (1964). Vampirism: A Review with New Observations. Archives of General Psychiatry 11(5):543–547.

Friday, July 29, 2022

Recent evolution in Estonia


Estonian women at a song festival (Wikicommons – Anastasia Lakhtikova)


Estonian women had more reproductive success during the late 20th century if they possessed a more masculine body build, narrower hips, and shorter legs. Such women married earlier and were less likely to stay on the mate market as long as possible.


Human evolution didn’t end in the Pleistocene. In fact, there has been more genetic change within our species over the past 10,000 years than over the previous 100,000, and perhaps more than over the previous million. The growing importance of culture did not slow down the pace of genetic change. In fact, culture became the main driving force of genetic evolution by replacing adaptation to a limited number of natural environments with adaptation to an ever-widening range of cultural environments (Cochran and Harpending 2009; Hawks et al. 2007; Rinaldi 2017).


Two years ago, I reviewed a study on recent evolution in the Estonian population (Frost 2020; Hõrak and Valge 2015). Among Estonians born between 1937 and 1962, women with only primary education had 0.5 to 0.75 more children than did women with tertiary education. This difference in reproductive success correlated with difference in cranial volume: children with larger crania were more likely to go on to secondary or tertiary education, independently of sex, socioeconomic position, and rural vs urban origin (Valge et al. 2019). Thus, for Estonian women in the late 20th century, higher education decreased fertility, probably by postponing the age of marriage.


That finding was found only for women. Perhaps Estonian men with higher education enjoyed greater reproductive success, in which case selection for less intelligent women may have been cancelled out by selection for more intelligent men.


The same research team has now published a new study of the same dataset, this time on both sexes. They confirm the original finding that female fertility correlated negatively with education and cranial volume. As for male fertility, although it correlated positively with education, the most fertile males had only average cranial volume. The authors had no explanation for that finding:


Stabilizing selection on the cranial volume of boys was an unexpected result, given that cranial volume in our study population predicts educational attainment independently of sex, socioeconomic background, and height. Since educational attainment was a strong predictor of fatherhood in our study, we would have expected positive directional selection on cranial volume. However, we found only evidence for stabilizing selection (Valge et al. 2022)


Perhaps women prefer men who are well-educated but not excessively intelligent. As one goes farther and farther away from the mean IQ of a population, higher intelligence becomes more and more often due to genetic “accidents”—unusual genetic variants or combinations of variants that may adversely affect other aspects of mind and behavior. A very intelligent person may seem autistic or have poor social skills.


The new study also shows that women had greater reproductive success if they possessed a more masculine body build, narrower hips, and shorter legs. That finding may seem counterintuitive. Don’t men prefer feminine-looking women? They do. However, as the authors show by citing earlier findings, shorter women are also less selective and likelier to marry earlier:


Similar reasoning might also explain why selection favored girls with masculine body build, narrow hips, and absolutely and relatively shorter legs in our study. If choosiness in women increases with desirability, this could lead to women with more feminine phenotypes engaging in a more time-consuming mate selection process, delaying their age of first birth, and thereby negatively affecting reproduction. (Valge et al. 2022)


Finally, the new Estonian study shows that heavier and stronger boys had more reproductive success.


The results relating to height and strength are consistent with studies of sexual selection showing that men who are taller, stronger, and more physically fit are generally perceived as more physically attractive by women, and therefore, have better opportunities for partnering and becoming a father. For instance, in a sample of Polish men born in the 1930s, childless men appeared significantly shorter than those with at least one child. In West Point graduates, the number of children increased linearly with height because taller men had higher probabilities of marrying more than once. Barclay and Kolk showed in a sample of 405,427 Swedish conscripts born between 1965 and 1972 that men in the lowest deciles of height, and in particular, physical fitness in early adulthood, had the lowest probabilities of transition to parenthood. (Valge et al. 2022)


Final thoughts


This is a study of Estonians who were born more than a half-century ago, long before the breakup of the Soviet Union. Things may be different now. Estonians have rapidly converged on Western social, behavioral, and ideological norms over the past three decades. Although their country is nominally independent, they are now strongly influenced by the inflow of Western culture via the media, and this new media environment is having a decisive impact on how they think and act (Karlin 2018).


Estonia is generally following the lead of the West. With respect to education and fertility, the negative correlation has become stronger throughout the West: “In all countries [Australia, United States, Norway, Sweden], however, education is negatively associated with childbearing across partnerships, and the differentials increased from the 1970s to the 2000s” (Thomson et al. 2014).


This differential is increasing not only between families but also within “families.” Second and third children are born increasingly to women who have divorced and are in relationships with low-quality fathers who often seem to be little more than sperm donors. In Norway, multi-partner fatherhood has become most common among men with the lowest level of education (10 years of schooling, "i.e., compulsory education"):


At age 45, about 15 percent of all men in the 1960-62 cohort with a compulsory education had had children with more than one woman, compared to about 5 percent among men with a tertiary degree. If looking at fathers only (Figure 6), the pattern becomes even more pronounced. At the lowest educational level, 19.3 percent of those who had become fathers, had children with more than one woman, compared to 6.1 percent of those at the highest educational level. (Lappegård et al. 2011)


This trend may partly explain the slowing down and reversal of the Flynn effect, i.e., the steady rise in mean IQ over the 20th century. There is some debate over whether the Flynn effect was a real increase in intelligence or simply an increase in familiarity with doing tests. In any case, its reversal seems real enough.


With respect to Norway, Bratsberg and Rogeberg (2018) have shown that the decline in mean IQ can be explained by “within-family variation.” In other words, mean IQ is declining among people who supposedly share the same genetic background, i.e., siblings. In Norway, however, siblings are increasingly half-siblings. Among Norwegian women with only two children, 13.4% have had them by more than one man. The figure rises to 24.9% among those with three children, 36.2% among those with four children, and 41.2% among those with five children (Thomson et al. 2014). 


The family unit is decomposing throughout the West. It is becoming little more than an administrative entity that can be repeatedly dissolved and reconstituted (Frost 2018a; Frost 2018b).





Bratsberg, B., and O. Rogeberg. (2018). Flynn effect and its reversal are both environmentally caused. Proceedings of the National Academy of Sciences 115 (26) 6674-6678


Cochran, G. and H. Harpending. (2009). The 10,000 Year Explosion: How Civilization Accelerated Human Evolution. Basic Books: New York.


Frost, P. (2018a). Why is IQ declining in Norway? Evo and Proud, June 19.


Frost, P. (2018b). Yes, the decline is genetic. Evo and Proud, June 26.


Frost, P. (2020). Declining intelligence in the 20th century: the case of Estonia. Evo and Proud, August 3.


Hawks, J., E.T. Wang, G.M. Cochran, H.C. Harpending, and R.K. Moyzis. (2007). Recent acceleration of human adaptive evolution. Proceedings of the National Academy of Sciences (USA) 104: 20753-20758.


Hõrak, P., and M. Valge. (2015). Why did children grow so well at hard times? The ultimate importance of pathogen control during puberty. Evolution, Medicine, and Public Health (1): 167–178,


Karlin, A. (2018). Gay marriage in Estonia. The Unz Review, October 30.


Lappegård, T., Rønsen, M., and Skrede, K. (2011). Fatherhood and fertility. Fathering 9: 103-120.


Rinaldi, A. (2017). We're on a road to nowhere. Culture and adaptation to the environment are driving human evolution, but the destination of this journey is unpredictable. EMBO reports 18: 2094-2100.


Thomson, E., T. Lappegård, M. Carlson, A. Evans, and E. Gray (2014). Childbearing across partnerships in Australia, the United States, Norway, and Sweden. Demography 51(2): 485-508. 


Valge, M., R. Meitern, and P. Hõrak.  (2022). Sexually antagonistic selection on educational attainment and body size in Estonian children. Annals of the New York Academy of Sciences Early view

Sunday, July 17, 2022

Cognitive evolution on the Italian Peninsula


A recent polygenic study has shown that mean cognitive ability is higher in the North of Italy than in the South. Cognitive evolution seems to have gone the farthest in the Northeast, perhaps because the Northwest earlier went through the Industrial Revolution, which severed reproductive success from economic success.




As a country, Italy came into existence only a century and a half ago. Regional differences are still strong, particularly between the North and the South. The “Southern question” is usually said to date from the unification of Italy in the 19th century:


In the decades following the unification of Italy, the northern regions of the country, Lombardy, Piedmont and Liguria in particular, began a process of industrialization and economic development while the southern regions remained behind. At the time of the unification of the country, there was a shortage of entrepreneurs in the south, with landowners who were often absent from their farms as they lived permanently in the city, leaving the management of their funds to managers, who were not encouraged by the owners to make the agricultural estates to the maximum. Landowners invested not in agricultural equipment, but in such things as low-risk state bonds. (Wikipedia 2022a)


De Rosa (1979) argues that the South had already fallen behind the North by the 18th century. At that time, its middle class was small, and economic relations were still structured by paternalism and familialism. One could go back even farther, to the Renaissance or even the late Middle Ages, to identify the moment when northern Italy, and Western Europe in general, embarked on sustained economic growth and thus pulled ahead of the rest of the world.


That sustained economic growth brought sustained demographic growth, particularly of the middle class. Gregory Clark found that the English middle class expanded steadily from the twelfth century onward, its descendants not only growing in number but also replacing the lower classes through downward mobility. By the 1800s, its lineages accounted for most of the English population. That demographic change coincided with mental and behavioral changes: higher cognitive ability, lower time preference, and a lower threshold for violent behavior. In a word, the English became more middle-class in character. “Thrift, prudence, negotiation, and hard work were becoming values for communities that previously had been spendthrift, impulsive, violent, and leisure loving” (Clark 2007, p. 166).


Elsewhere in Western Europe, the middle class similarly expanded during late medieval and early modern times. The result would be a growing contrast between regions that had participated in this economic and demographic change and those that had not, such as southern Italy. The contrast can be seen not only on purely economic measures but also on mental ones, like the INVALSI standardized test—an annual test of skills in Italian schools. It is divided into two sections: Italian language skills and Math skills. On both tests, northern Italian students do better than southern Italian students, the difference being a little over half a standard deviation:


Yes, the North-South gap in academic achievement could have a purely environmental cause—and this is a recurring problem when we try to tease apart genetic and cultural evolution. If economic development is held back by a culture of poverty, that same culture may discourage students from trying to do better at school. Those students may also have less access to proper nutrition, medical care, libraries, and so on.


Polygenic scores for cognitive ability


That is why there is so much interest in measures of innate cognitive ability. The most promising one is the polygenic score (PGS)—the summation of alleles (genetic variants) that have been associated with cognitive ability, as measured by educational attainment.  At present, we have identified enough of these alleles to explain 11-13% of the overall variation in cognitive ability (Lee et al. 2018).


Yes, those alleles are just a sample of the total number, but why would they be an unrepresentative sample? More to the point: why would PGS data show certain geographic patterns and not a lot of random noise? The mean PGS does indeed differ geographically among human populations. It is highest in Eurasia, with East Asians, Ashkenazi Jews, and Finns having the highest scores. That geographic pattern is in line with IQ data (Piffer 2019).


Polygenic scores on the Italian Peninsula


In a recent study, Piffer and Lynn (2022) have found regional differences in Italy for alleles associated with educational attainment. They used two datasets: one encompassing 129 Italian individuals and the other 947. All of these individuals had all four grandparents born in the same part of Italy (this requirement was imposed to eliminate the effects of recent interregional migration). When the authors grouped the data into three large regions—North, Central, and South—they found “a clear north-south gradient, with central Italians occupying an intermediate position.” There was more overlap between central and southern Italians than between central and northern Italians.


The datasets were too small to show genetic differences within each of the three large regions. If we go back to the INVALSI data, we see that academic achievement is much stronger in the North-Northeast (Lombardia, Trentino, Veneto, Friuli) than in the Northwest (Valle d’Aosta, Liguria).


At first thought, that geographic pattern may seem counter-intuitive. In northern Italy, industrialization began in the northwest and came later to the northeast: “the diffusion of industrialisation that characterised the northwestern area of the country largely excluded Venetia and, especially, the South” (Wikipedia 2022b). If economic development had driven cognitive evolution on the Italian Peninsula, why would this evolution have gone farther in the northeast? Why would it be negatively associated with industrialization?


Because the Industrial Revolution put a stop to cognitive evolution. It severed the link between economic success and reproductive success. Previously, businesses were family-run, and the family provided the workforce. Successful business owners were incentivized to have larger families, and their children would have the means to marry at a younger age. Then, in the late 19th century, that stage of economic development began to give way to industrial capitalism. Financial success no longer translated into early marriage and large families who could help with the work. If more workers were needed, they would simply be hired. Business owners now tended to have smaller families because of the high maintenance costs of middle-class children (Canlorbe and Frost 2020; Frost 2018).


Cognitive evolution thus ended earlier in the Northwest of Italy than in the Northeast. By the same token, interregional migration has had more time to erode the cognitive advantage that evolved in the Northwest. Yes, the datasets were limited to people who had all four grandparents born in the region, but, for most people, that limitation would not eliminate the effects of interregional migration before the mid-20th century.





Canlorbe, G., and P. Frost (2020). Why are human groups so different? American Renaissance, March 20.  


Clark, G. (2007). A Farewell to Alms. A Brief Economic History of the World, 1st ed. Princeton University Press: Princeton, NJ, USA.


De Rosa, L. (1979). Property Rights, Institutional Change, and Economic Growth in Southern Italy in the XVIIIth and XIXth Centuries. Journal of European Economic History 8(3): 531-551.


Frost, P. (2018). Rise of the West. Part II. Evo and Proud, December 27  


Lee, J. J., Wedow, R., Okbay, A., Kong, E., Maghzian, O., Zacher, et al. (2018). Gene discovery and polygenic prediction from a genome-wide association study of educational attainment in 1.1 million individuals. Nature Genetics 50(8): 1112-1121.


Piffer, D. (2019). Evidence for Recent Polygenic Selection on Educational Attainment and Intelligence Inferred from Gwas Hits: A Replication of Previous Findings Using Recent Data. Psych 1(1): 55-75.    


Piffer, D., & Lynn, R. (2022). In Italy, North-South Differences in Student Performance Are Mirrored by Differences in Polygenic Scores for Educational Attainment. Mankind Quarterly 62(4), Article 2.  


Wikipedia (2022a). Economy of Italy – Southern Question  


Wikipedia (2022b). Economic history of Italy.  


Thursday, July 7, 2022

Adapting to bubonic plague


Rash associated with familial Mediterranean fever (Wikicommons – Dr. H.J. Lachman)


In the eastern Mediterranean, people were likelier to survive bubonic plague if they had stronger inflammatory responses to infection in their lungs, gut, and other tissues. Today, that natural selection is attested by a high incidence of familial Mediterranean fever.


Familial Mediterranean fever is due to mutations that increase the body’s production of pyrin, a protein that assists inflammatory responses to infection of the lungs, gut, and other tissues. It’s common among eastern Mediterranean peoples, like Jews, Syrians, Armenians, Turks, Greeks, and Italians. The most common symptoms are inflammation of the abdominal lining, the joints, and the chest. Some kind of natural selection seems likely because different mutations have evolved independently to produce the same disease within the same geographic region.


Several years ago, Greg Cochran suggested this fever might be an adaptation to trypanosome parasites (Leishmaniasis). Recent research now suggests an adaptation to bacteria of the genus Yersinia. Y. pestis causes bubonic plague, whereas Y. pseudotuberculosis and Y. enterocolitica cause gastroenteritis. These pathogens are highly infectious because they decrease the body’s production of pyrin and thus reduce its ability to fight infection. To compensate for this underproduction of pyrin, there seems to have been selection for mutations that cause overproduction of pyrin, hence the high incidence of familial Mediterranean fever in populations that have coexisted with Yersinia bacteria (Loeven et al. 2020).


Yepiskoposyan and Harutyunyan (2007) argue that selection for familial Mediterranean fever must have begun more than 2,500 years ago, since one of the alleles responsible for it is found in different communities of the Jewish diaspora, notably Iraqi and North African Jews. That argument doesn’t convince me, since diaspora communities were not reproductively isolated. The mutation could have arisen in one community and then been spread to others by Jewish individuals moving from one place to another.


I’m inclined to believe that selection for this fever began with the earliest recorded outbreak of bubonic plague: the Plague of Justinian (541-549 AD), which killed an estimated 25 million people throughout the Mediterranean Basin and the Middle East. An earlier date is nonetheless possible, since Y. pestis has been attested in archaeological finds as far back as 5,000 years ago (Wikipedia 2022).





Chung, L.K., Y.H. Park, Y. Zheng, I.E. Brodsky, P. Hearing, D.L. Kastner, J.J. Chae, and J.B. Bliska. (2016). The Yersinia Virulence Factor YopM Hijacks Host Kinases to Inhibit Type III Effector-Triggered Activation of the Pyrin Inflammasome. Cell Host Microbe 20(3):296-306.


Cochran, G. (2015). Familial Mediterranean fever. West Hunter, January 9.


Loeven, N.A., N.P. Medici, and J.B. Bliska. (2020). The pyrin inflammasome in host-microbe interactions. Curr Opin Microbiol 54:77-86.


Wikipedia (2022). Bubonic plague.


Yepiskoposyan, L., and A. Harutyunyan. (2007) Population genetics of familial Mediterranean fever: a review. Eur J Hum Genet 15: 911–916.

Thursday, June 30, 2022

Extreme founder events among ancestral Europeans



Expansion of steppe pastoralists (Narasimhan et al. 2019)


How much of the present-day European gene pool comes from the indigenous hunter-gatherers? How much from Neolithic farmers of Anatolian origin? And how much from steppe pastoralists of the Don-Volga area? There is no easy answer.



A “founder event” results when a few people split off from their original population and found a new one. The fewer they are, the likelier they will differ genetically, on average, from the original population. The founder event is a “bottleneck” through which only a small fraction of the original genetic diversity can pass into the new population.


A founder event is one of three reasons why adjacent populations may differ from each other genetically. The other two are:


Natural selection – The boundary between two adjacent populations often corresponds to a change in the natural environment (vegetation, climate, etc.) or the cultural environment (diet, way of life, rules and prohibitions, sexual division of labor, etc.). The two populations are thus subjected to different regimes of natural selection.


Population admixture or replacement – One of the two adjacent populations has admixed with or been replaced by a population that has moved into the area.


In practice, a founder event overlaps with differences in natural selection. When pioneers move into a new area, they tend to be better suited to the local conditions than the people they left behind. The less suited are less likely to go, and if they do go they are more likely to go back … or die. So, from the outset, there already is some selection.


A new study of ancient DNA has shown that founder effects have been more frequent and more “extreme” than previously thought: “In humans, we find that over half of the analyzed populations have evidence for recent founder events, associated with geographic isolation, modes of sustenance, or cultural practices such as endogamy” (Tournebize et al. 2022, p. 1)


Contrary to popular belief, Ashkenazi Jews are not the outcome of a particularly extreme founder event:


Across worldwide populations, we identified 53 groups that have experienced more extreme founder events (with significantly higher founder intensity) than AJs [Ashkenazi Jews], who have high rates of recessive diseases due to their history of founder events (Tournebize et al. 2022, p. 7)


Perhaps those recessive diseases are not due to a founder event. Perhaps they are a side-effect of selection for an adaptive trait. If a founder effect had been the cause, those diseases would be distributed randomly over different metabolic pathways. Actually, they are all associated with excessive storage of sphingolipids, a key component of neural tissue (Cochran et al. 2006; Diamond 1994).


Founder events seem to have been frequent among ancestral Europeans, regardless of whether they were hunter-gatherers, farmers, or pastoralists. But those events seem to have been more “intense” among European hunter-gatherers. In other words, founder groups were smaller and spent more time passing through the population bottleneck.


Recent analysis has shown that present-day Europeans are a mixture of three major ancestry groups related to ancient European hunter-gatherers, Anatolian farmers, and Eurasian Steppe pastoralists. […] Across the three groups, we found that the frequency of founder events was similar, ranging between 90–100%. However, the average founder intensity was significantly higher in European hunter-gatherers […] compared to the Near Eastern farmers […] or the Steppe pastoralists. (Tournebize et al. 2022, p. 11)


Hunter-gatherers had more intense founder events because they had a lower population density. Founder groups were thus smaller and less representative of the original population from which they came.


… we found local hunter-gatherer groups had more extreme founder events than the Neolithic farmers or Bronze Age individuals. This suggests that population sizes in Europe have increased over time, coupled with changes in ancestry and transitions in lifestyle. Our results are consistent with a recent study that measured short runs of homozygosity in ancient Europeans and found a similar increase in population size during the Neolithic period. Our results are also in agreement with archeological evidence for increased population size during the Neolithic transition. (Tournebize et al. 2022, p. 14)


This takes us back to the previous quote: Europeans are a mix of indigenous hunter-gatherers, Anatolian farmers, and steppe pastoralists. These three groups, and their roles in European prehistory, can be summarized as follows:


·         Farmers began to move into Europe from present-day Turkey about 10,000 years ago. Initially, they advanced rapidly through territory inhabited by small nomadic bands.

·         About 7,000 years ago, the wave of advance stalled along a line running from the Low Countries to the Black Sea. To the north, along the North Sea and the Baltic, were large semi-sedentary communities of hunter-fisher-gatherers who could less easily be replaced because they were so numerous.

·         About a thousand years later, farming resumed its northward advance, although the advance was now much more a matter of people adopting farming rather than being replaced by farmers.

·         Meanwhile, around 5,400 years ago, some hunter-gatherers in the Don-Volga area adopted pastoralism and began to expand westward into Europe and southeastward into the Middle East, Central Asia, and South Asia. They may have been ancestral Indo-Europeans.


How much did each of the three groups contribute to the European gene pool? Which group contributed the most and which the least? The question is hard to answer, for three reasons:


Double counting


The hunter-gatherers of Mesolithic Europe contributed to the present European gene pool both directly and indirectly. The steppe pastoralists were themselves indigenous hunter-gatherers who had adopted pastoralism, plus an admixture of up to 18% from Anatolian farmers. As the Anatolian farmers pushed into Europe, they became gradually “Europeanized” through intermixture with local hunter-gatherers. It is also possible that the Anatolian farmers were themselves the product of an earlier expansion of European hunter-gatherers into the Middle East (Frost 2014).


Founder events


We measure population replacement by measuring the degree of genetic difference between the original group and the one that replaced it. Is that a valid method? Let’s take the replacement of hunter-gatherers by farmers, and let’s assume that all of the farmers were descended from hunter-gatherers who had adopted farming. The two groups would still be genetically different. The farmers would have been the product of a founder event—a small and genetically unrepresentative group of hunter-gatherers who had decided to take up farming.


Differences in natural selection


Population replacement is hard to measure for another reason: hunter-gatherers and farmers lived under different regimes of natural selection. They were selected for their ability to adapt to different diets, types of shelter, and means of subsistence. To go from one way of life to the other required not only cultural change but also genetic change.


For instance, the population frequency of haplogroup U shows a sharp break at the time boundary between late hunter-gatherers and early farmers (Bramanti et al. 2009). That break strongly suggests that the original Europeans were largely replaced by farmers spreading into Europe from the Middle East. Yet haplogroup U would persist in Denmark at high frequencies long after the transition to farming (Melchior et al. 2010). In Latvia and Ukraine it would persist into Neolithic times (Jones et al. 2017). Haplogroup U probably disappeared from the European gene pool because it ceased to be adaptive. It has been shown to shift the energy balance away from ATP synthesis and toward production of body heat, a useful adaptation if you sleep in makeshift shelters and pursue game in all kinds of weather (Balloux et al. 2009; Montiel-Sosa et al. 2006). It’s less useful if you sleep in a warmer environment and can plan your outdoor activities.




Whenever I make this argument, the counter-argument is that founder events and natural selection could not possibly explain all of the genetic difference we see between late hunter-gatherers and early farmers in Europe. I agree. I’m just saying that the magnitude of the demographic replacement has been overestimated.




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Tournebize, R., G. Chu, and P. Moorjani. (2022). Reconstructing the history of founder events using genome-wide patterns of allele sharing across individuals. PLoS Genet 18(6): e1010243.

Thursday, June 23, 2022

Ancestral East Asians and adaptation to coronaviruses


Early farming village in China (Wikicommons – Xinyang City Museum, Gary Todd)


Respiratory viruses began to propagate more easily when hunting and gathering gave way to farming and as settlements grew larger. Humans may have then evolved to use coronaviruses as a natural vaccine against deadlier respiratory diseases, like tuberculosis and pneumonia.



A new genomic study has found that East Asians had to adapt to epidemics of coronaviruses some 25,000 years ago. The authors looked at gene variants for proteins that interact with coronaviruses in five East Asian populations: Han Chinese (Beijing); Han Chinese (South China); Dai (Yunnan, China); Japanese; and Vietnamese. There were three main findings:


·         Ancestral East Asians had to adapt to coronavirus epidemics around 25,000 years ago

·         They adapted by acquiring mutations that are close to genes that regulate the development of lung tissue and other tissues affected by COVID-19

·         Those mutations either promote or block infection by coronaviruses (Souilmi et al. 2021, p. 3505).


The last finding is puzzling. Did those ancestral East Asians become more vulnerable or less vulnerable to coronaviruses? The authors simply say that half of those mutations from 25,000 years ago have “anti- or proviral effects” versus 29% of all proteins that interact with coronaviruses (Souilmi et al. 2021, p. 3509). Fine. But how many of those mutations were antiviral and how many proviral?


It might seem strange that natural selection would actually make people more susceptible to coronavirus infections. Yet such susceptibility could be beneficial. A viral infection can boost immunity to other pathogens, including deadly ones that cause tuberculosis, pneumonia, or pneumonic plague. Until recently, coronaviruses were typically mild in their effects, producing what we call the “common cold.” They may thus act as a natural vaccine against deadlier respiratory diseases (Frost 2020).


Respiratory diseases are believed to have become serious for humans when hunting and gathering gave way to farming. People became sedentary and their settlements grew larger with time, thus providing respiratory viruses with better conditions for propagation (Comas et al. 2013). This theoretical model is in conflict, however, with the above finding that ancestral East Asians began adapting to coronaviruses some 25,000 years ago, long before they adopted farming and became sedentary. We’re thus left with the unlikely conclusion that coronavirus epidemics began among scattered bands of hunter-gatherers.


The estimate of 25,000 years ago is probably wrong. The authors arrived at that figure by calculating the latest date when the ancestors of the four East Asian groups were still a single population. But East Asians are not descended from a single population. Their origins are best described by the "Two-Layer" (TL) hypothesis:


·         Modern humans spread into East Asia through a northern route and a southern route.

·         The southerners were then replaced to varying degrees by northerners who spread out of northeast Asia and successively occupied northern China, southern China, and Southeast Asia (Oxenham and Buckley 2016; Xu et al. 2006).

·         Thus, as you go farther south in East Asia, the population has a greater admixture from the earlier southern “layer”—from hunter-gatherers who closely resemble the relic groups that still exist in parts of Southeast Asia, i.e., the Andaman Islanders, the Aeta of the Philippines and the Maniq and Semang of the Malayan Peninsula.


Admixture from that older southern substrate pushes back in time the latest common ancestors, who never existed. Adaptation to coronaviruses therefore happened at a later date, probably when the “northerners” pushed into what is now northern China and adopted farming. They then grew in population, pushed farther south, and intermixed with the hunter-gatherers who lived there. 





Comas, I., M. Coscolla, T. Luo, et al. (2013). Out-of-Africa migration and Neolithic coexpansion of Mycobacterium tuberculosis with modern humans. Nature Genetics 45: 1176–1182.


Frost, P. (2020). Does a commensal relationship exist between coronaviruses and some human populations? Journal of Molecular Genetics 3(2): 1-2.


Frost, P. (2022). A natural vaccine. Evo and Proud, February 21


Oxenham, M., and H.R. Buckley. (2016). The population history of mainland and island Southeast Asia, in M. Oxenham and H.R. Buckley (eds) The Routledge Handbook of Bioarchaeology in Southeast Asia and the Pacific Islands. Routledge.


Souilmi, Y., M.E. Lauterbur, R. Tobler, C.D. Huber, A.S. Johar, S.V. Moradi, W.A. Johnston, N.J. Krogan, K. Alexandrov, and D. Enard. (2021). An ancient viral epidemic involving host coronavirus interacting genes more than 20,000 years ago in East Asia. Current Biology 31(16), 3504–3514.e9.


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