Tuesday, August 05, 2008

Male dominance not all that?   posted by Razib @ 8/05/2008 02:54:00 PM
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Tyler points me to a new paper coming out in PNAS, Male dominance rarely skews the frequency distribution of Y chromosome haplotypes in human populations. It isn't on the site yet, but New Scientist has a write up:
To determine whether dominance could last more than a couple generations, Watkins and a team of anthropologists and geneticists sifted through the DNA of 1269 males from 41 Indonesian communities.
...
Out of 41 communities, from Bali to Borneo to mainland Indonesia, only five showed evidence of long-term dominance by a few male lines.
...
Of course, Genghis Khan proves that some powerful males can ensure their lineage - if not through prosperity, then promiscuity - but such men are rare, Watkins says.

"If I were to take 100 random Mongolians and follow their family lines, I wouldn't have seen anything special."


How does Wilkins know this about Mongolians? Perhaps there's some empirical data in the paper he isn't reporting re: Mongolia, but it seems that one must be cautious about extrapolating from Indonesia. As most of you know, Indonesia is an archipelago, and water tends to be really good at bottling up gene flow.

Of course, this is a big question that spans all human societies across time. I assume there are going to be variations across space, and time, and that frequency dependence is important as a conditional which frames any assertions we make. I suspect that Genghiside "super-male" lineages are more a feature of the last 10,000 years where it is possible for only a few people to sequester large amounts of surplus productivity and travel was much more common along elites. Additionally, I've made this point before, but I'll do so again: in many pre-modern societies being a high status male opens you up to a great deal of risk and gain simultaneously. So there might be a long term angle in turning your vehicles into indispensable betas....

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Monday, June 02, 2008

Selection speculation: CLOCK and reward-dependence in Africans   posted by agnostic @ 6/02/2008 02:39:00 AM
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Since so many comments lead off with some variant of "I would guess," why not try to corrall them all into one post where they could serve a purpose? Each week I'll find some area of the human genome that shows signs of recent selection, see what phenotypes the gene affects, and although I'll likely provide the most convincing story, readers can conjecture to their heart's content about what might have driven selection. It may, for once, improve the discussion to comment while still deranged from last night's drink.

Let's start with the data: using Haplotter, we see that for the gene CLOCK, there is a signal of recent selection in Africans but not in Europeans or Asians. The CLOCK gene is involved in maintaining our circadian rhythm, and I started this search looking for between-group differences in being "day people" vs. "night people." However, other genes related to circadian rhythm -- PER1, PER2, PER3, Tim, CSNK1E, Cry1, and Cry2 -- show no signals of recent selection anywhere. So perhaps there is something else that CLOCK does that these others do not.

It turns out that CLOCK is also involved in the dopaminergic system, which regulates mood and behavior. None of the OMIM entries for the other circadian rhythm genes mentions dopamine, mood, the reward system, etc. So it is more likely that CLOCK has undergone selection for its effects on mood and on the brain's reward system, rather than on circadian rhythm per se. In mice, and so potentially in humans as well, mutations in CLOCK make individuals lower in anxiety and higher in risk-taking (Roybal et al. 2007; free full text).

Turning to data from Lynn & Martin (1995), which I turned into a convenient graph here, we see that Nigerians -- the African group that HapMap data come from -- are indeed the lowest in the world in the personality trait Neuroticism (the Israeli data-point must be a mis-coding by Lynn, who is known for doing so). They are also the highest in the world in Extraversion. Neuroticism measures a tendency toward anxiety and related states, while Extraversion measures, among other things, pace of living (manics living a faster-paced life) and excitement-seeking. So Nigerians show a psychological and behavioral phenotype that's fairly comparable to the mice with mutations in CLOCK, strengthening the hypothesis that selection at CLOCK has acted on personality rather than circadian rhythm per se.

Now comes the fun part -- telling a story about why this phenotype was more advantageous in Western Africa than in Central Europe or Northeast Asia over the past 10,000 or so years. The low-anxiety and high-excitement-seeking phenotype is suspiciously like that associated with derived alleles at the DRD4 locus, and the novelty-seeking 7R allele at that locus does show up at intermediate frequencies in Africans (Chen et al. 1999; see a map here). I concur with Harpending & Cochran (2002) (free full text), who argued in the DRD4 case that the phenotype is more advantageous in "cad societies" than in "dad societies."

More concretely, I suggest, based on the teachings of the pickup artist community, that low anxiety aids in approaching more females in a shorter time -- you are less paralyzed during the approach, and you get over rejection more easily -- and that higher risk-taking motivates you to approach more females. Contingent upon having other traits that are appealing to females (a muscular body, artistic skill, dancing or singing ability, a knack for flirting, and so on), the phenotype here would have increased male success in courtship of (multiple) females, a stronger determinant of reproductive success in a "cad society" than in a "dad society."

Lastly, here are two testable predictions: 1) lowland South American and highland New Guinean tribes should also show signs of selection at the CLOCK locus, since they are even more prototypically "cad societies," and they have the highest frequencies of the 7R allele at DRD4. And 2) African Bushmen should show no signs of selection at CLOCK since they are more "dad societies" and have lower frequencies of the 7R allele at DRD4. (ALFRED does not have data on CLOCK.)

The fact that Northeast Asians do not show signs of selection at CLOCK already supports the hypothesis, since they are more "dad societies" and have a very low frequency of 7R at DRD4 (and even the 2R allele, which shows up somewhat there, has a much more muted effect than does 7R). The lack of selection at CLOCK in Central Europeans is neither here nor there: they are more "dad societies" than Western Africa but not so much as East Asia; Europeans also have intermediate frequencies of 7R and DRD4. So we could have an example of "Rushton's Rule," where East Asians show no selection for low-anxiety / high-novelty-seeking alleles at either DRD4 or CLOCK, Europeans show selection at just one locus, and West Africans show selection at both.

To the best of my knowledge, as they say, this is the first argument for why Africans show a signal of selection at CLOCK. Consider it part of the graduate student with a PC trend in studying human evolution. You heard it here first.

References

Chen, C., M. Burton, E. Greenberger, & J. Dmitrieva (1999). Population migration and the variation of Dopamine D4 receptor (DRD4) allele frequencies around the globe. Evolution and Human Behavior, 20(5): 309-324.

Harpending, H. & G. Cochran (2002). In our genes. PNAS, 99(1): 10-12.

Lynn, R., & T. Martin (1995). National differences for thirty-seven nations in extraversion, neuroticism, psychoticism and economic, demographic and other correlates. Personality and Individual Differences, 19(3): 403-406.

Roybal, K., D. Theobold, A. Graham, J.A. DiNieri, S.J. Russo, V. Krishnan, S. Chakravarty, J. Peevey, N. Oehrlein, S. Birnbaum, M.H. Vitaterna, P. Orsulak, J.S. Takahashi, E.J. Nestler, W.A. Carlezon, Jr., & C.A. McClung (2007). Mania-like behavior induced by disruption of CLOCK. PNAS, 104(15): 6406-11.

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Tuesday, January 01, 2008

Why are models tall?   posted by Razib @ 1/01/2008 12:22:00 AM
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First, happy New Year! Second, when I saw this post from Matt in the referrals I assumed he was going to be offering some skepticism as to the utility of formal models in science. No, not really, not that type of model. The sort that our resident callipygiaphile ruminates upon.

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Sunday, December 30, 2007

Malthus, innovation's friend?   posted by Razib @ 12/30/2007 12:43:00 PM
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Since the previous post was about the tendency toward radical skepticism and subjectivism within cultural anthropology, I thought I would point to this piece in The Economist which highlights positive insights from various anthropological fields. The article emphasizes the possible role that population pressure and the quest for food might have had in spurring human innovation, from the atlatl to agriculture. An interesting point to note is the implicit suggestion that high rates of hunter-gatherer warfare might have constrained population pressure and possibly lead to relatively higher standards of living; something familiar from Greg Clark's model. From a population genetic angle, I am curious as to whether the endemic warfare of cultures which were pre-state resulted in higher or lower Nm*?

* N = population, m = migration rate. Nm > 1 results in equilibration of allele frequencies across demes, while Nm < 1 tends to lead to divergence.

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Tuesday, October 02, 2007

Inferences about patrilocality from genetics   posted by Razib @ 10/02/2007 11:03:00 PM
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When genetics is used as a supplement to history or anthropology it should ideally offer more precision and reduce the field of possibilities. Unfortunately that is not always the case. For example, when asking big questions such as "Are humans 'naturally' patrilocal?" genetics has generally come up with conflicting results. The fact is that about 70% of "traditional" societies are patrilocal; wives move to live with husbands and their family. Some genetic evidence seems to support this (e.g., greater mtDNA [female] variation than Y [male] lineage variation), but there is also variation within populations and some skepticism as to whether such simple comparisons between different uniparental loci really can give a definitive answer.

This paper in PLOS ONE tackles the question using comparisons with chimpazees to help calibrate our expectations, The Genetic Signature of Sex-Biased Migration in Patrilocal Chimpanzees and Humans:
...Here we review some methodological reasons for these inconsistencies, and take them into account to provide an unbiased characterization of mtDNA and NRY variation in chimpanzees, one of the few mammalian taxa where males routinely remain in and females typically disperse from their natal groups. We show that patterns of mtDNA and NRY variation are more strongly contrasting in patrilocal chimpanzees compared with patrilocal human societies. The chimpanzee data we present here thus provide a valuable comparative benchmark of the patterns of mtDNA and NRY variation to be expected in a society with extremely female-biased dispersal.



Homo Sapiens, are a complex species. We're capable of a lot and our level ofsocial complexity isn't truly cognitively tractable for the mental toolkit that evolution has given us. So we make recourse to generalizations and first-order approximations. The problem with this is that sometimes the deviation away from the central tendency is as interesting and evolutionarily salient as the mode on the frequency distribution of the trait. When we say that 70% of human societies are patrilocal, that means that 30% are not. That also doesn't mean that 70% of human societies have always been patrilocal. It also doesn't quantify how much deviation there is from social expectation, e.g., perahps in the 70% of societies which are patrilocal a substantial minority of males move to the locality of the female, or, males and females from the same locality marry each other so that the whole dichotomy is rendered irrelevant (though even if you live within a village whether you reside in the relatives of the female or relatives of the male in an extended family circumstance is also an important point). Generalizing about the modal behavior of our species may leave the concept more analytically tractable, but it may also render our model inaccurate toward the point of uselessness.

On the other hand, chimpanzees aren't that complex...at least compared to humans. Yes, I know that chimps are the geniuses of the animal kingdom, but the chimp is the Peter Keating of the animal world and we are the Howard Roarks. Unlike humans with their facultative complexity chimpanzee societies are pretty uniform. While there are exceptions to the rule (as in the famous Gombe troop), female chimps tend to leave their natal group while males tend to remain in their natal group. In shot, chimps seem to be much more obligate in their patrilocality than humans. The workers above note that both chimp lineages, the bonobo and common chimp, are patrilocal, so this is likely an ancestral characteristic of the clade dating back at least 1 million years. It seems that chimpanzees exhibit a much more unambiguous genetic signature of patrilocality than humans, the ratio of Y to mtDNA variance is significantly lower than in humans.

There are some technical reasons why the human results could exhibit problems. For example, mtDNA might simply be more diverse than Y lineages for endogenous reasons (higher mutational rates?). Or there might be lower effective population size of one sex, which would skew the variation of the uniparental lineages independent of deme-to-deme gene flow. There are also issues of coarseness of analysis, the authors point out that in some human communities Fst analyses were preformed by spanning tribal groups where intermarriage was rather uncommon, while ignoring the pervasiveness of between deme gene flow on the intra-tribe level. Finally, there is the issue that human societies change. The Japanese were once matrilocal, but now the are patrilocal. Within historic times many matrilineal societies have shifted toward patrilocal practices (though sometimes there are ghosts of matrilineal practice, in ancient Egypt marriage to a woman of the royal line was often essential to solidify the claims of the male claimant). There variations over time can obscure or erase genetic patterns and replace them with new ones, and periodic oscillations would presumably result in a meta-stable level of diversity which balances out both mtDNA and Y.

Chimps can get around some of these issues. Chimpanzee societies are more homogeneous in their behavior patterns, have been studied for nearly two centuries, and have been characterized to a very fine level of demic structure in terms of their social dynamics (even to the point of familial histories). In short, chimpanzees are empirically tractable because of their small numbers and limited set of behaviors. The similarity across the two clades and their relative homogeneity should reassure one that temporal variation is minimized. Because of their charismatic nature chimpanzees have also been tracked in a way which makes assessments of male vs. female migration patterns on the level of specific demes tenable (i.e., you couldn't get funding to do such detailed research on most species, so people have to engage in a lot more guesswork in terms of how they behave when no one is looking, which can explain why ethologists were long fooled by "monogamous" birds before DNA fingerprinting cleared up some issues).

It seems only the most extremely patrilocal and polygynous human societies in the sample approached the chimp norm. To me this suggests that we'll have to be a bit more careful and qualified when talking about human patrilocality. Though we're not totally malleable and subject to great constraints, our behavioral flexibility is orders of magnitude more developed than the faculties of our chimp cousins. It shouldn't surprise if we tend to explore a far greater sample space of social systems, not only because we can, but also because we inhabit so many environmental and cultural ecosystems. Polyandry which consists of brothers marrying one woman in Tibet did not arise because it was natural; rather, it seems to be the best opportunity for reproduction that a low status male could attain in that society where resources were at a premium. It was a functional response to a specific set of circumstances. This work focuses on the dimension of male and female migration across adjacent demes, classical models of gene flow. And this likely works well for chimpanzees, and perhaps hunter-gatherers. But it seems excessively oversimplified when it comes to the mass societies which arose after the rise of agriculture. 4,000 years ago a man who was born in Switzerland was buried at Stonehenge. In their day the Mongol hordes swept from the Pacific to the plains of Pannonia, from Baikal to Baghdad. This was certainly a scattering of Y lineages of immense scale. These sorts of movements don't really work well with chimpanzee analogs...because, well, chimps have never produced transcontinental hordes! Atop the simmering activity of deme-to-deme wife and husband exchanges there will periodically flash a fire of migratory activity. Sometimes this will be a total folk movement, but I suspect that more often what you would see was a migration of males. There is much more variation in human social networks than among chimpanzees, so we should focus less on the mean outcome as opposed to the structure and pattern of the variation and its distribution.

Note: You can discuss this paper over at PLOS ONE if you so choose!

Langergraber KE, Siedel H, Mitani JC, Wrangham RW, Reynolds V, et al. (2007) The Genetic Signature of Sex-Biased Migration in Patrilocal Chimpanzees and Humans. PLoS ONE 2(10): e973. doi:10.1371/journal.pone.0000973

Related: Sperm competition.

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Thursday, September 13, 2007

Four Stone Earth @ John Hawks   posted by Razib @ 9/13/2007 05:22:00 PM
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Four Stone Hearth, the anthropology blog carnival, is over at John Hawks' place.

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Friday, May 11, 2007

Tasmanian Aboriginal DNA not extracted...   posted by Razib @ 5/11/2007 05:28:00 PM
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Here is a story about a museum which returned Tasmanian Aboriginal remains to a community group without doing DNA analysis, because they argued that this sort of compromise would preserve them for future analysis. This is a tricky area. I'm obviously generally disinclined to sympathize with "communities" who claim bones because the individuals making the decisions are simply individuals, and often individuals playing politics. That being said, the history of the Tasmanian Aboriginals was characterized by proactive bestial treatment on the part of Europeans, and the relationship of the native peoples of Australasia and "civilization" has been highly "problematic." I think that from a cost vs. benefit standpoint targeting descendants of Aboriginal groups is perhaps a better bet, community groups can't make a strong argument that they have a right to decide whether someone gives genetic material or not. I'd honestly be curious to hear what John Hawks thinks, seeing as he's an anthropologist and so has presumably thought this issue through to some extent.

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Tuesday, April 17, 2007

The wild man of the forest   posted by Razib @ 4/17/2007 10:26:00 AM
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Since we're talking about chimps, check out this long story about the species which surveys the opinions and research of a range of primatologists.

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Saturday, April 07, 2007

Just don't use the word 'race'   posted by sustaSe @ 4/07/2007 03:35:00 PM
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That seems to be the conclusion of a recent review of Nick Wade's Before the Dawn published in Nature Genetics:
My reluctance to recommend the book stems also from Wade's discussions of 'race' and biology. I agree with Wade that there is something biological about racial categories. In my opinion, although racial identity is socially negotiated, people use physical traits as cues when 'assigning' a racial identity to themselves or another individual. Racial categorization isn't blind to biology. ... Although one might detect biological differences between races, any highlighting of the racial categories (just a subset of groups with biological correlates) has social costs, according to recent social science research. On the other hand, Neil Risch, cited often in the book, has argued that there are significant (medical) costs of ignoring the relationship between racial categories and biology. I suggest that these different costs be weighed in each circumstance where one might link 'race' and genetics. Wade's broad description of races as clearly delineated biological entities is unjustified in the context of a book about human history intended for a general audience. Why use the term 'race', when 'geographic ancestry' or 'continental origin' are more accurate and less costly in social terms, especially since Wade's definition of 'race' is "continent of origin"? I suggest acknowledging the correlation between racial labels and continents of origin, and saving the term 'race' for contexts in which social costs are outweighed by other costs.


This is not the argument I expected to follow the sentence "My reluctance to recommend the book stems also from Wade's discussions of 'race' and biology." In this case, the author isn't being snide by putting race in quotes, as she really means the word race rather than its referent. How often do scholars write that consternation over race is largely related to extra-scientific concerns?

However, I have to criticize this argument, at least to the extent that I'm able to examine the evidence presented. A footnote to the "recent social science research" showing that using the word 'race' is harmful (but that cryptic synonyms are OK) would be appreciated, as this forms the basis of the argument against discussing 'race'. Is it only harmful to discuss 'race' and 'genetics' or 'biology'? Is the attribution of racial differences to environmental/cultural causes not similarly harmful? Is it really true, as is implied, that Wade is morally obliged to substitute most instances of "race" in his text with "continent of origin"?

There's a lot to commend in this review, largely stemming from the reviewers' honesty and directness, and especially in contrast with this hatchet job published in the sister journal Nature.

Update - full text:
Given the rich content of Nicholas Wade's latest book, Before the Dawn, I wish I could simply recommend the book, describe its highlights and stop there. Wade provides a valuable overview of the last ten years of scientific literature on genetic insights into the history of our species. He is an excellent storyteller, weaving the scientific results into a thrilling tale of human migration and settlement, competition and warfare, cultural and linguistic evolution and environmental challenges. The history of our species is a fascinating one, and Wade brings it to life.

I congratulate Wade for taking great pains to qualify many of his statements with terms such as "seems" and "appears to." In an important, related vein, early in the book he notes that any "intent" suggested in biologists' statements about evolution reflects shorthand communication and is not meant to imply that evolution has any particular goal "in mind." Evolutionary biologists will certainly appreciate that note. Furthermore, given that few readers will be specialists in all the fields represented in the book (paleoanthropology, archaeology, linguistics, genetics and more), many will appreciate Wade's practice of defining terms.

Despite the book's many strengths, I am reluctant to recommend the book unconditionally. I found some sections of the book challenging to read, as I looked for supporting evidence for various claims. For example, Wade suggests that the San, peoples in southern Africa who subsist via foraging, are the "closest living approximation to the ancestral human population." Behaviorally, this might be true. However, Wade goes on to suggest that the San may not have evolved genetically, as "foragers have presumably had much the same environment for the last 50,000 years." Wade appears to be unaware of the diverse environments even today within sub-Saharan Africa; furthermore, the changing global climate over the past 50,000 years has often had dramatic impacts on humans living in Africa.

Although at many points in the book Wade notes the speculative nature of conclusions from genetic, archaeological or geographic data, he occasionally treats those conclusions as fact elsewhere. For example, he writes, "There is no way to know for certain the nature of the interaction between the two human species [anatomically modern humans and Neanderthals]." Yet elsewhere he writes, "...[the Neanderthals] crushed the attempt by anatomically modern humans to penetrate the Levant." The reader is at risk of being lulled by numerous "maybes," "seems" and "appears" into trusting unsupported but confidently stated comments elsewhere in the book.

My reluctance to recommend the book stems also from Wade's discussions of 'race' and biology. I agree with Wade that there is something biological about racial categories. In my opinion, although racial identity is socially negotiated, people use physical traits as cues when 'assigning' a racial identity to themselves or another individual. Racial categorization isn't blind to biology. Yet Wade puts words in the mouths of the American Anthropological Association (AAA) when he states that the AAA "dismisses the idea that biological differences can be recognized between races." He backs up his statement with an AAA quote that makes a different point: "any attempt to establish lines of division among biological populations [is] both arbitrary and subjective." Although one might detect biological differences between races, any highlighting of the racial categories (just a subset of groups with biological correlates) has social costs, according to recent social science research. On the other hand, Neil Risch, cited often in the book, has argued that there are significant (medical) costs of ignoring the relationship between racial categories and biology. I suggest that these different costs be weighed in each circumstance where one might link 'race' and genetics. Wade's broad description of races as clearly delineated biological entities is unjustified in the context of a book about human history intended for a general audience. Why use the term 'race', when 'geographic ancestry' or 'continental origin' are more accurate and less costly in social terms, especially since Wade's definition of 'race' is "continent of origin"? I suggest acknowledging the correlation between racial labels and continents of origin, and saving the term 'race' for contexts in which social costs are outweighed by other costs.

Wade's chapter on language is replete with details of relationships among languages, methodology for reconstructing those relationships and arguments in support of methods that are purported to give ages of languages. Although much of this discussion will undoubtedly provoke many linguists, the most provocative element in this chapter is a more general statement: "The mutability of language reflects the dark truth that humans evolved in a savage and dangerous world in which the deadliest threat came from other human groups." I see little support for this conjecture. Language, at least a language rich in elements, cannot come into being without being mutable. And as Wade notes earlier in the book, "Language would have made small groups more cohesive, enabled long-range planning and fostered the transmission of local knowledge and learned skills." Mutability may reflect these advantages rather than a "savage and dangerous world."

Where I am familiar with the relevant scientific literature, I see the details that Wade includes in this, his latest book, as accurately representing scientific findings. Wade often wraps these scientific details in dramatic stories, thereby creating a book both informative and entertaining. However, some of Wade's general themes, such as his claim of a very high level of aggressiveness of prehistoric hunter-gatherers, are just that—dramatic stories. Readers will benefit most by considering each such claim as one among several plausible interpretations of the data.

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