Showing posts with label classes. Show all posts
Showing posts with label classes. Show all posts

Saturday, July 4, 2020

Recent evolution of the British population



Frederick Morgan – Off for the Honeymoon (Wikicommons) Over the past 2,000 years, the British gene pool has shifted toward alleles that favor lighter hair, sunburn, and educational attainment. Was this because high-status men tended to mate with blonder, fairer women?



Have we evolved over the past two thousand years? Until recently, the answer was thought to be 'no.' Cultural evolution took over from genetic evolution around the time farming took over from hunting and gathering, some ten thousand years ago, thus putting our ancestors on a path to increasing social complexity: sedentary living, growth of towns and villages, formation of states, trade and specialization of labor, and so on. It was culture that changed during recorded history, not genes.

Well, things are not that simple. Genes and culture have coevolved with each other. Yes, culture has been changing rapidly over the past ten thousand years. But so have genes. During that time, our genetic evolution has been driven by adaptation not only to natural environments but also to cultural environments. Increasingly so. We live more and more in cultural environments of our making (Chen et al., 2016; Cochran and Harpending 2009; Hawks et al. 2007).

In what ways have we changed genetically during the past ten thousand years? In the ways we digest food. With the shift to dairy farming, and the resulting increase in milk consumption by adults, natural selection favored those who could digest milk sugar, an ability previously confined to infants.

We have also changed in the ways we think and behave. That kind of evolution is not difficult. A few point mutations may alter a behavior by changing its timing, its intensity, or its threshold of stimulation. Other alterations have been much more polygenic. Cognitive ability, for instance, seems to have increased through mutations at many genes, with each mutation causing only a tiny fraction of the increase.

Because recent evolutionary change has so often been polygenic, we need to examine it in relation to many genetic variants spread over the entire genome, i.e., by means of genome-wide association studies. Such studies can take many forms. A recent one, proposed by Stern et al. (2020), may be better than earlier versions, particularly in avoiding biases due to population structure and population stratification.

I nonetheless have a few reservation about this proposed method:

1. Population stratification can be a factor in evolutionary change. Let's take the work of Gregory Clark on the growth of the English middle class. He found it grew 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. Parallel to that demographic growth, English society became more and more middle class in its values. "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). Isn't that evolutionary change through population stratification? Or am I missing something?

2. The new method can reveal only evidence of directional selection. It thus fails to capture other interesting forms of selection, like diversifying selection.


How the British have evolved over the past 2,000 years

Stern et al. (2020) used their method to study how the British population has evolved over the past two thousand years. They found increases in the prevalence of lighter hair, in tanning and sunburn, in age at first birth, in bone mineral density, and in the risk of type 2 diabetes. They also found decreases in the risk of neuroticism and in the risk of high glycated hemoglobin levels.

Some of these changes correlate with each other. In such cases, we should step back and try to identify the common cause.

Lighter hair, more sunburn ... and higher educational attainment

Over the past 2,000 years, the British gene pool has shifted toward alleles that favor lighter hair, sunburn, and educational attainment. These changes in allele frequency correlate with each other, so what, exactly, was driving the overall change?

There is genetic linkage between light hair and pale skin, but it's weak. In fact, pale skin often coexists with dark hair. Moreover, we still have to explain the link to educational attainment. The common cause for all three changes may have been sexual selection mediated by social class. In other words, high-status men tended to mate with blonder, fairer women.

This form of sexual selection was observed in a Japanese study on social class and skin color. Upper-class men were shown to be fairer-skinned than lower-class men, even when the latter were factory workers and not farmers and even though the measurements were taken on unexposed skin. Wealthier men have a wider range of prospective brides and can thus choose the fairest women, for "skin color has long been regarded, by the Japanese, as one of the criteria for evaluating physical attractiveness, especially in young females" (Hulse 1967). Similarly, in India "[w]ealthy landowning families often have a tradition of seeking light-skinned brides among poorer members of their subcaste. It is very common to find a high concentration of lighter-skinned people among established land-owning families" (Béteille 1967).

Darwin discussed this sexual selection with reference to English social classes:

Many persons are convinced, as it appears to me with justice, that our aristocracy, including under this term all wealthy families in which primogeniture has long prevailed, from having chosen during many generations from all classes the more beautiful women as their wives, have become handsomer, according to the European standard, than the middle classes; yet the middle classes are placed under equally favorable conditions of life for the perfect development of the body. (Darwin 1936[1888], p. 892)

Until the 20th century, higher social status meant higher fertility (Clark 2007). Thus, the physical and mental characteristics of the upper and middle classes tended to displace those of the lower class.

Higher risk of Type 2 diabetes and glycated hemoglobin

Why would natural selection favor type 2 diabetes? Isn't diabetes harmful? It is, in a modern environment that lets you ingest calories almost without limit. That wasn't the case in Britain for most of the past two thousand years. During that time, food was scarce for most people, and natural selection favored the ability to get as many calories as possible out of our food.

Less neuroticism

This evolutionary change may be related to the demographic success of the middle class and associated mental and behavioral traits, particularly lower time preference and higher future orientation. The nascent English middle class valued being “calm, cool, and collected,” as opposed to reacting emotionally to negative outcomes.


References

Béteille, A. (1967). Race and descent as social categories in India. Daedalus 96(2): 444-463.

Chen, C., R.K. Moyzis, X. Lei, C. Chen, and Q. Dong. (2016). The encultured genome: Molecular evidence for recent divergent evolution in human neurotransmitter genes. In: J.Y. Chiao, S.-C. Li, R. Seligman, and R. Turner, Eds, The Oxford handbook of cultural neuroscience. New York, NY: Oxford University Press, 315-336.

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

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

Darwin, C. (1936 [1888]). The Descent of Man and Selection in relation to Sex. reprint of 2nd edition, The Modern Library, New York: Random House.

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, 104(52), 20753-20758.

Hulse, F.S. (1967). Selection for skin color among the Japanese. American Journal of Physical Anthropology 27(2): 143-156.

Hysi, P.G., A.M. Valdes, F. Liu, N.A. Furlotte, D.M. Evans, V. Bataille, et al. (2018). Genome-wide association meta-analysis of individuals of European ancestry identifies new loci explaining a substantial fraction of hair color variation and heritability. Nature Genetics 50(5): 652-656.

Morgan, M.D., E. Pairo-Castineira, K. Rawlik, O. Canela-Xandri, J. Rees, D. Sims, A. Tenesa, and I.J. Jackson. (2018). Genome-wide study of hair colour in UK Biobank explains most of the SNP heritability. Nature Communications 9: 5271

Neel, J. V. (1962). Diabetes mellitus: a 'thrifty' genotype rendered detrimental by 'progress'? American Journal of Human Genetics 14: 353-362.

Stern, A.J., L. Speidel, N.A. Zaitlen, and R. Nielsen. (2020). Disentangling selection on genetically correlated polygenic traits using whole-genome genealogies
bioRxiv 2020.05.07.083402

Saturday, January 21, 2012

Do classes become castes?


Relative frequencies of surnames of the rich and the poor (common criminals) 1236-1858. In England, there seems to have been much downward mobility among descendants of the medieval rich and some upward mobility among descendants of the medieval poor (Clark, 2010).
H/T to Jason Malloy


Henry Harpending (2012) argues that a meritocracy would become a caste society in a few generations:

Consider free meritocracy in a two-class system, meaning that for each generation anyone in the lower class who has greater merit than someone in the upper class immediately swaps class with them. Mating then occurs at random within class.

[…] Class mobility after the first generation is 30% while after four generations it has declined to 10% and continues to decline after that. The average merit in the two classes is about -1SD in the lower and +1SD in the upper on the original scale, corresponding to IQs of 85 and 115.

[…] after four generations, about 70% of the variance is between classes.

This model, however, contradicts what Clark (2009a) found in his historical study of surnames and social class in England. He first collected rare English surnames that were exclusive in the year 1600 to the rich (as represented by wealthy testators) or to the poor (as represented by common criminals). He then went forward in time to the year 1851 and determined the occupational profile of the same rare surnames:

How do the descendants of these two groups look in terms of socioeconomic status by 1851? Surprisingly there seems to be almost complete regression to the mean.

Between 1600 and 1851, there was apparently great downward mobility among descendants of the rich and modest upward mobility among descendants of the poor. Clark (2010) subsequently found that this regression held true for the entire period stretching from 1236 to 1858 (see above chart).

Why does this outcome diverge so much from the theoretical outcome described above? One reason is the assumption that marriage takes place only within each social class. Yet assortative mating is only a tendency, and exceptions are numerous. In post-medieval England, a widower would likely take a second wife of lower social status because of his disadvantaged position on the marriage market, given the care required for his existing children. It was also accepted for an upper-class man to marry a woman of lower rank, on the condition that she be beautiful. This phenomenon was noted by Darwin:

Many persons are convinced, as it appears to me with justice, that our aristocracy, including under this term all wealthy families in which primogeniture has long prevailed, from having chosen during many generations from all classes the more beautiful women as their wives, have become handsomer, according to the European standard, than the middle classes; yet the middle classes are placed under equally favourable conditions of life for the perfect development of the body. (Darwin, 1936 [1888], p. 892)

Another reason is downward mobility. In England, the upper and middle classes were reproductively more successful than the lower class until the late 19th century. But higher-class families could offer their children only a limited number of occupational slots. A certain proportion thus had to emigrate or move down the social ladder. The lower class was thereby continually replenished by the demographic overflow of the upper and middle classes.

Finally, the word “merit” has different meanings in different contexts. It is never just IQ. In post-medieval England, merit meant a mix of “middle-class” values: thrift, self-control, future time orientation, and rejection of violence as a way to settle disputes (Clark, 2007; Clark, 2009a; Clark, 2009b). In other societies, merit may involve a different mix of predispositions and personality traits, such as ruthlessness and willingness to use violence.

Yet caste societies do exist. How do they come about? The main precondition seems to be not only the existence of social classes, but also a monopoly on certain occupations by each class. In such circumstances, downwardly mobile individuals cannot compete with the existing lower class, since the latter’s livelihood remains off-limits.

Take Japan. That country had a social evolution similar to England’s, i.e., gradual demographic expansion of the middle class and, correspondingly, gradual demographic replacement of the lower classes by downwardly mobile individuals. But the lowest class, the Burakumin, survived because it had a monopoly on occupations that involved taking life or handling dead bodies (e.g., leather working, butchery, undertaking, etc.). The Burakumin thus survive as a remnant of the majority Japanese population that existed several centuries ago.

Stigmatized castes, like the Burakumin, may provide a window into a population’s evolutionary past. Such groups cannot participate in the gene-culture co-evolution of the majority population. Nor do they have much leeway for their own gene-culture co-evolution, since they are rigorously confined to a few occupations.

References

Clark, G. (2007). A Farewell to Alms. A Brief Economic History of the World. Princeton and Oxford: Princeton University Press.

Clark, G. (2009a). The indicted and the wealthy: surnames, reproductive success, genetic selection and social class in pre-industrial England.
http://www.econ.ucdavis.edu/faculty/gclark/Farewell%20to%20Alms/Clark%20-Surnames.pdf

Clark, G. (2009b). The domestication of Man: The social implications of Darwin. ArtefaCTos 2(1): 64-80.

Clark, G. (2010). Regression to mediocrity? Surnames and social mobility in England, 1200-2009
http://www.econ.ucdavis.edu/faculty/gclark/papers/Ruling%20Class%20-%20EJS%20version.pdf

Darwin, C. (1936) [1888]. The Descent of Man and Selection in relation to Sex. reprint of 2nd ed., The Modern Library, New York: Random House.

Harpending, H. (2012). Class, Caste, and Genes, West Hunter, January 13
http://westhunt.wordpress.com/2012/01/13/class-caste-and-genes/