Wednesday, August 6, 2008

Aleksandr Solzhenitsyn RIP

Aleksandr Solzhenitsyn passed away on Monday. He was one of several writers who have truly touched me (George Parkin Grant was another). His main message, for us in the West, is that Communism was not specific to Russian culture and history. This message is summarized by Wikipedia:

It is a popular view that the October revolution of 1917 resulting in a violent totalitarian regime was closely connected to Russia's earlier history of tsarism and culture, especially that of Ivan the Terrible and Peter the Great. Solzhenitsyn claims this is fundamentally wrong and famously denounced the work of Richard Pipes as "the Polish version of Russian history". Solzhenitsyn argues Tsarist Russia did not have the same violent tendencies as the Soviet Union. For instance, in Solzhenitsyn's view, Imperial Russia did not practise censorship; political prisoners were not forced into labour camps and the number of political prisoners was only one ten-thousandth of those in the Soviet Union; the Tsar's secret service was only present in the three largest cities, and not at all in the army. The violence of the Communist regime was in no way comparable to the lesser violence of the Tsars.

He considered it far-fetched to blame the catastrophes of the 20th century on one 16th century and one 18th century czar, when there were many other examples of violence which could have inspired the Bolshevik in other countries earlier in time, especially mentioning similarities with the Jacobins of the Reign of Terror of France.

Communism was a kind of idealism that allowed no dissent. Some debate did exist, but only within narrowly circumscribed bounds. Is abstract art progressive? What is the timetable for building socialism? And so on. There was no substantive debate over core values.

With no debate, there could be no true reform. There could, at best, be tinkering. And such tinkering often made things worse. Eventually, even the Soviet elite realized that fundamental changes were needed. But how?

Imagine a machine with no ‘off’ switch. Imagine a machine that does not hesitate to destroy its own creators. Imagine a machine that can be stopped only by the limitations of reality itself. That was Russia’s nightmare.

And it could become ours. I used to think that such a thing could happen only in Russia. Now, looking around my own country, I must confess I was wrong. It could happen here. And it could happen for the noblest of intentions.

"In our country the lie has become not just a moral category but a pillar of the State."
–Aleksandr Solzhenitsyn, December 29, 1974

Thursday, July 31, 2008

Lewontin's Fallacy?


Turning to race, we must begin with the fraught question of whether it even exists, or whether it is instead a social construct. The Harvard geneticist Richard Lewontin originated the idea of race as a social construct in 1972, arguing that the genetic differences across races were so trivial that no scientist working exclusively with genetic data would sort people into blacks, whites, or Asians. In his words, "racial classification is now seen to be of virtually no genetic or taxonomic significance."

Lewontin's position, which quickly became a tenet of political correctness, carried with it a potential means of being falsified. If he was correct, then a statistical analysis of genetic markers would not produce clusters corresponding to common racial labels.

In the last few years, that test has become feasible, and now we know that Lewontin was wrong. Several analyses have confirmed the genetic reality of group identities going under the label of race or ethnicity. In the most recent, published this year, all but five of the 3,636 subjects fell into the cluster of genetic markers corresponding to their self-identified ethnic group. When a statistical procedure, blind to physical characteristics and working exclusively with genetic information, classifies 99.9 percent of the individuals in a large sample in the same way they classify themselves, it is hard to argue that race is imaginary.

The above is from an article by Charles Murray in Commentary, “The Inequality Taboo.” It apparently refers to an earlier article on ‘Lewontin’s Fallacy’ by Edwards (2003).

Murray is right in believing that human genetic variation does cluster geographically and that these clusters are adaptively significant —be they ‘races’, ‘geographical populations’ or whatever.

But is this point proven by the above line of reasoning? Lewontin never argued that human genetic variation is random. He simply affirmed that human races, however they may be defined, account for only a small percentage of total variation. Hence, there is far more variability within than between human populations. Murray counters that this conclusion is false because Lewontin looked at only one genetic trait at a time.

Clearly, if two groups overlap, they are more easily told apart with several criteria than with just one. If we use enough criteria, the overlap will shrink to zero: individuals will be assignable to either group with no ambiguity. But none of this means that within-group variability has decreased. In fact, it has actually increased. The only difference now is that this variability consists of combinations of genes that are unique to each group. How does this fact invalidate Lewontin’s contention that “the largest part by far of human variation [is] accounted for by the differences between individuals.”?

One might object that Charles Murray was talking about genes that contribute to intelligence and that such a contribution is almost certainly polygenic. Yes, but we’re not looking at several genes that display one form in one group and another form in the other. The two groups are still very heterogeneous whether you’re looking at any one gene or at gene combinations.

To find the flaw in Lewontin’s argument, we must examine his initial assumption: a random sample of genes should tell us how important race differences are. True, a large enough sample of genes will tell us whether a species has begun to differentiate into identifiable subpopulations. It will also tell us, roughly, when these subpopulations began to differentiate from each other.

But it won’t tell us how important between-population differences are in relation to within-population differences. It’s an apples and oranges comparison. The two groups of genes are qualitatively different.

First, when genes vary between populations, it’s usually because these populations inhabit different environments with different sets of selection pressures. Genes that differ across this environmental boundary are necessarily genes that make a difference, i.e., that have selective value.

In contrast, when genes vary within a population, despite similar selection pressures, it’s usually because they have little or no selective value (or because they form a balanced polymorphism, but that’s another topic!).

Second, the genetic markers used by population geneticists (blood groups, enzymes, mtDNA, etc.) tend to be selectively less important. This is partly because that is how population geneticists want them to be. Researchers will often choose markers that are close to selective neutrality because such markers change at a predictable rate (through random mutations) and can thus provide a time clock of a population’s history.

Such markers are also chosen because their protein products are easier to find and measure in body tissues. These ‘structural proteins’ are usually similar when we compare different species or even different genera. Humans and chimps, for instance, look very much alike when it comes to the protein building blocks that make up their body tissues. We have diverged from other apes largely through evolutionary changes at a higher level. i.e., regulatory genes that control development and other higher-order processes.

This point was grasped by Stephen J. Gould (1977, 406). He explained how we distort our understanding of genetic variation by relying on data from structural genes:

The most important event in evolutionary biology during the past decade has been the development of electrophoretic techniques for the routine measurement of genetic variation in natural populations. Yet this imposing edifice of new data and interpretation rests upon the shaky foundation of its concentration on structural genes alone (faute de mieux, to be sure; it is notoriously difficult to measure differences in genes that vary only in the timing and amount of their products in ontogeny, while genes that code for stable proteins are easily assessed).

References

Edwards, A.W.F. (2003). Human genetic diversity: Lewontin’s fallacy. BioEssays, 25, 798-801.

Gould, S.J. (1977). Ontogeny and Phylogeny. Belknap Press: Cambridge (Mass.)

Lewontin, R.C. (1972). The apportionment of human diversity. Evolutionary Biology, 6,381-398.

Murray, C. (2005). The inequality taboo. Commentary, September.

Tuesday, June 17, 2008

More genetic variation within than between?

Research in human genetics has highlighted that there is more genetic variation within than between human groups, where those groups are defined in terms of linguistic, geographic, and cultural boundaries.

Statement 2. Guiding principles on using racial categories in human genetics (Soo-Jin Lee et al., 2008)

The above statement is part of an open letter to human geneticists, recently published in Genome Biology. The authors are a number of Stanford University scholars from the humanities, social sciences, life sciences, law and medicine.

Yes, the statement is true. It has been confirmed by a wide range of studies, the most well known being by Richard Lewontin (1972). At most human genes, there is far more variation within human populations than between them. Lewontin concluded that 85% of human genetic variation exists only between individuals and cannot be apportioned on a population basis. In this, he was repeating a conclusion that others had made before him, notably Frank Livingstone (1962), and that others still have since confirmed. Clearly, racial categories seem to be very hazy entities—so hazy, in fact, as to seem useless.

Or maybe not. It turns out that the same haziness exists between many species. When we compare pairs of species that are closely related but anatomically distinct, we often see more genetic variation within each species than between them, so much so, that we cannot reliably assign individuals to either by genetic criteria alone (see older post).

How come? When two populations diverge and eventually become separate species, they usually do so because they have adapted to different environments with different selection pressures. These selection pressures, however, do not act on the entire genome. In fact, they act only on a small set of genes—the ones that need to function differently in either environment.

We see the same thing with artificial selection. Kennel clubs maintain distinct dog breeds by insisting that dogs in any one breed meet various criteria. This distinctness disappears, however, when we look at their genomes.

… genetic and biochemical methods … have shown domestic dogs to be virtually identical in many respects to other members of the genus. … Greater mtDNA differences appeared within the single breeds of Doberman pinscher or poodle than between dogs and wolves. Eighteen breeds, which included dachshunds, dingoes, and Great Danes, shared a common haplotype and were no closer to wolves than poodles and bulldogs. These data make wolves resemble another breed of dog.

… there is less mtDNA difference between dogs, wolves, and coyotes than there is between the various ethnic groups of human beings, which are recognized as a single species.
(Coppinger & Schneider, 1995)

Artificial selection uses a smaller set of criteria than does natural selection. Nonetheless, both forms of selection act on only a small fraction of the genome, partly because most genes are of marginal selective value and partly because many genes can function identically in a wide variety of species.

Of course, even at these other genes two populations may start to show diverging patterns of variation once they have become reproductive isolated species. But this is a long process that occurs through each of them slowly and separately accumulating its own mutations at genes of low selective value. This is not the case with our species. We began to spread out of Africa only 50,000 or so years ago.

References

Coppinger, R. and R. Schneider (1995). Evolution of working dogs. In J. Serpell (ed.), The Domestic Dog: Its Evolution, Behaviour and Interactions with People. Cambridge: Cambridge University Press, pp. 21-47.

Soo-Jin Lee, S., Mountain, J., Koenig, B., Altman, R., Brown, M., Camarillo, A., Cavalli-Sforza, L., Cho, M., Eberhardt, J., Feldman, M., Ford, R.,Greely, H., King, R., Markus, H., Satz, D., Snipp, M., Steele, C., and Underhill, P. (2008). The ethics of characterizing difference: guiding principles on using racial categories in human genetics. Genome Biology, 9, 404 doi:10.1186/gb-2008-9-7-404

Lewontin, R.C. (1972). The apportionment of human diversity. Evolutionary Biology, 6,
381-398.

Livingstone, F.B. (1962). On the non-existence of human races. Current Anthropology, 3, 279-281.

Women's hair color. It doesn't pay to be average

Gene expression has run a second post on blonde preference. This time the analysis is based on the front covers of Maxim magazine, whose readership is 27.5 years old on average and thus provides a better guide of what young men prefer (Playboy’s readership is 32.5 on average).

The post makes two points:

1) After increasing from the mid-1960s to about the year 2000, preference for blonde women seems to have levelled off over the past 11 years.

2) In comparison to the white American population, there is an overrepresentation of women with either blonde or dark brunette hair on the front covers of Maxim. Women with intermediate hair shades are underrepresented.

The second finding is interesting because it bears out the frequency-dependent nature of hair-color preference. Men seem to have a stronger preference for less common shades.

The Rise of the Blonde Playmate

Gene Expression has a post on changing preferences for hair color over time. The hair color of Playboy playmates was charted from 1954 to 2007, on the assumption that this monthly series reflects male sexual tastes. It turns out that the percentage of blonde playmates has risen over the years: from a low of about 35% in the mid-1960s to a high of 60% around the year 2000. A similar study was done fifteen years ago, with similar results (Rich & Cash, 1993).

These proportions are well above the actual proportion of blonds among white Americans. When Rich and Cash (1993) studied a sample of undergraduates, the proportions were 68.1% brunette, 26.8% blond, and 5.1% red. This breakdown parallels those put forward in two British studies: 68% brunette, 25% blond, 1% red, and 6% black (Takeda et al., 2006); and 74% brunette; 18% blond, and 8% red (Mather et al., unpublished). Jason Malloy says a similar breakdown appears in Beddoe (1885).

Thus, blond hair preference does not reflect the actual distribution of hair color. In fact, in light of current demographic trends, the scarcer blondes become, the more they seem to be preferred.

This frequency dependence has been shown in humans. Thelen (1983) presented male participants with slides showing attractive brunettes and blondes and asked them to choose, for each series, the woman they would most like to marry. One series had equal numbers of brunettes and blondes, a second 1 brunette for every 5 blondes, and a third 1 brunette for every 11 blondes. Result: the rarer the same brunette was in a series, the likelier the men would choose her.

Just as blondes are more strongly preferred as they become scarcer, they seem to be less popular when more abundant. Havelock Ellis (1928, pp. 182-183) noted a weaker preference for blonde women in England than in France, which he ascribed to the higher prevalence of blondness among the English.

Indeed, if this preference were not frequency-dependent, sexual selection should have steadily increased the incidence of blond hair, to the point of eliminating all other colors. Yet blonds do not form the majority of any human population. Even Swedes are no more than 40% blond, and this is using a definition that already includes a variety of shades (platinum blond, ash blond, sunny blond, sandy blond, golden blond, strawberry blond, zebra blond, dirty blond, brownish blond, see Wikipedia article).

In addition to America’s changing demographics, the rise of the blonde playmate may also reflect the spread of Playboy magazine into American Catholic communities where it was initially excluded and where blond hair is less common (many of these communities being of southern European origin). In the wake of Vatican II (1962-1965), Catholic Americans have adopted increasingly secular attitudes towards sex, and consumption of soft porn has become less stigmatized, if not tolerated.

References

Beddoe, J. (1885). The Races of Britain: A Contribution to the Anthropology of Western Europe, Arrowsmith, Bristol & Trübnermm, London.

Ellis, H. (1928). Studies in the Psychology of Sex. Vol. IV, "Sexual Selection in Man." Philadelphia: F.A. Davis Company.

Mather, F., Manning, J.T., & Bundred, P.E. (unpublished). 2nd to 4th digit ratio, hair and eye colour in Caucasians: Evidence for blond hair as a correlate of high prenatal oestrogen.

Rich, M.K., & Cash, T.F. (1993). The American image of beauty: Media representations of hair color for four decades. Sex Roles, 29, 113-124.

Takeda, M.B., Helms, M.M., & Romanova, N. (2006). Hair color stereotyping and CEO selection in the United Kingdom. Journal of human behavior in the social environment, 13, 85-99

Thelen, T.H. (1983). Minority type human mate preference. Social Biology, 30, 162-180.

The Big Bang of modern human evolution

The Smithsonian magazine has an interesting article on the origin of modern humans and the events leading to their rapid spread ‘out of Africa’:

Then, about 80,000 years ago, says Blombos archaeologist Henshilwood, modern humans entered a "dynamic period" of innovation. The evidence comes from such South African cave sites as Blombos, Klasies River, Diepkloof and Sibudu. In addition to the ocher carving, the Blombos Cave yielded perforated ornamental shell beads—among the world's first known jewelry. Pieces of inscribed ostrich eggshell turned up at Diepkloof. Hafted points at Sibudu and elsewhere hint that the moderns of southern Africa used throwing spears and arrows. Fine-grained stone needed for careful workmanship had been transported from up to 18 miles away, which suggests they had some sort of trade. Bones at several South African sites showed that humans were killing eland, springbok and even seals. At Klasies River, traces of burned vegetation suggest that the ancient hunter-gatherers may have figured out that by clearing land, they could encourage quicker growth of edible roots and tubers. The sophisticated bone tool and stoneworking technologies at these sites were all from roughly the same time period—between 75,000 and 55,000 years ago.

The above archaeological findings match African mtDNA data presented by Watson et al. (1997):

We find that most African mitochondrial sequences appear to be the result of demographic expansions that started ~60,000-80,000 years ago, the earliest of which led to the colonization of Eurasia. Only a minority (13%) of sequences fall outside these expansion clusters, echoing a time, before the expansions, when the human mitochondrial gene pool was possibly more diverse (in terms of mean sequence divergence) than it is today.

These population expansions appear to have been driven by cultural innovations that gave some African populations an edge over others and, eventually, over archaic humans in Europe and Asia. Unfortunately, instead of speaking of an expansion out of Africa, the Smithsonian article prefers the term ‘exodus’ — as if life had become so intolerable in Africa that humans were forced to pick up and leave. In fact, almost the opposite happened: modern humans did so well in Africa that they elbowed out their archaic rivals not only on their own continent but on others as well.

The Smithsonian article also speculates about the reasons behind their demographic success. These humans had become better at constructing mental models of the world around themselves. They could imagine the things they had to make and tinker with them in their minds. And they could share this mental tinkering with others through language.

This improvement in mental functioning may have been made possible initially by increased consumption of fatty acids in seafood. Then, as humans developed a new cultural environment and its attendant demands on brainpower, there would have been selection for genes to ‘hardwire’ this neurological change, i.e., the Baldwin effect.

Virtually all of these sites had piles of seashells. Together with the much older evidence from the cave at Pinnacle Point, the shells suggest that seafood may have served as a nutritional trigger at a crucial point in human history, providing the fatty acids that modern humans needed to fuel their outsize brains: "This is the evolutionary driving force," says University of Cape Town archaeologist John Parkington. "It is sucking people into being more cognitively aware, faster-wired, faster-brained, smarter." Stanford University paleoanthropologist Richard Klein has long argued that a genetic mutation at roughly this point in human history provoked a sudden increase in brainpower, perhaps linked to the onset of speech.

References

Gugliotta, G. (2008). The Great Human Migration. Why humans left their African homeland 80,000 years ago to colonize the world. Smithsonian magazine, July.

Watson, E., Forster, P., Richards, M., and Bandelt, H-J. (1997). Mitochondrial footprints of human expansions in Africa. American Journal of Human Genetics, 61, 691-704.

Maps of European hair and eye color

A reader sent me the following e-mail:

I want to ask you some questions regarding your maps.

To be honest both maps look very questionable. Also I didn't see any source for these studies, especially the one for eye color looks completely invented. I'm sure that the percentage of light eyes in Italy (for example) is higher than 1-19%, surely in Italy there are more people with light eyes than Spain, Portugal and especially than Turkey or Lebanon , as there are more people with light eyes in Southern England than Bosnia.

My suppositions are confirmed by the studies of Coon or Lundman for example:

Map by Carleton Coon

Map by Bertil Lundman


The maps on my website come from my article on European hair and eye color diversity (Frost 2006) and are reproduced from an anthropology textbook (Beals & Hoijer 1965, pp. 213-214). Beals and Hoijer, in turn, cite a textbook by another anthropologist, Frederick Hulse (1963: p. 328). Unfortunately, Hulse does not indicate the provenance of his data. I suspect he was using data from military recruits, with a lot of interpolation. Or perhaps he was using even earlier maps.

Since my 2006 article came out, I have found one such map in a work by Biasutti (1959, cap. I Nazioni europee e la loro composizione etnica p. 43), which in turn is taken from Günther (1929). At Gene Expression, Razib has turned up another map by Carleton Coon (1982), which is ascribed to ‘Elmer Rising, 1939.’

All of these maps resemble each other in their broad outlines, but the minor differences are significant. I still have not found the original data on which any of them are based.

References

Beals, R.L. and H. Hoijer. (1965). An Introduction to Anthropology, 3rd edition, New York: MacMillan Co.

Biasutti, R. (1959). Razze e Popoli della Terra. Torino: Unione Tipografico-Editrice

Coon, C.S. (1982). Racial Adaptations. Burnham.

Coon, C.S. (1939). The Races of Europe. New York: The Macmillan Co.

Frost, P. (2006). European hair and eye color - A case of frequency-dependent sexual selection? Evolution and Human Behavior, 27, 85-103.

Günther, H.F.K. (1929). Rassenkunde des deutschen Volkes. Munich: J.F. Lehmann

Hulse F.S. (1963). The Human Species. An Introduction to Physical Anthropology. New York: Random House.

Lundman, B.J. (1977). The Races and Peoples of Europe. New York: IAAEE