Sunday, October 05, 2008

South Park   posted by Razib @ 9:59 PM
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New episode of South Park this Wednesday. As always you can watch it online.


news.thinkgene.com   posted by Razib @ 7:08 AM
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Think Gene has a new digg-like site up, news.thinkgene.com. Worth checking out.

Friday, October 03, 2008

Beyond Belief III   posted by Razib @ 7:01 AM
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Beyond Belief III is going down right now, the 3rd to the 6th. Free registration is soldout, but it looks like there are still some pay slots. If you're in the San Diego area you might be able to attend....

Wednesday, October 01, 2008

Russian dudes are imperialists   posted by Razib @ 5:23 AM
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Dynamics of Alliance Formation and the Egalitarian Revolution:
Background:

Arguably the most influential force in human history is the formation of social coalitions and alliances (i.e., long-lasting coalitions) and their impact on individual power. Understanding the dynamics of alliance formation and its consequences for biological, social, and cultural evolution is a formidable theoretical challenge. In most great ape species, coalitions occur at individual and group levels and among both kin and non-kin. Nonetheless, ape societies remain essentially hierarchical, and coalitions rarely weaken social inequality. In contrast, human hunter-gatherers show a remarkable tendency to egalitarianism, and human coalitions and alliances occur not only among individuals and groups, but also among groups of groups. These observations suggest that the evolutionary dynamics of human coalitions can only be understood in the context of social networks and cognitive evolution.


Their conclusion:

We propose a simple and flexible theoretical approach for studying the dynamics of alliance emergence applicable where game-theoretic methods are not practical. Our approach is both scalable and expandable. It is scalable in that it can be generalized to larger groups, or groups of groups. It is expandable in that it allows for inclusion of additional factors such as behavioral, genetic, social, and cultural features. Our results suggest that a rapid transition from a hierarchical society of great apes to an egalitarian society of hunter-gatherers (often referred to as “egalitarian revolution”) could indeed follow an increase in human cognitive abilities. The establishment of stable group-wide egalitarian alliances creates conditions promoting the origin of cultural norms favoring the group interests over those of individuals.


The title is a joke because the lead author is Sergey Gavrilets, who is a product of Moscow State University in the 1970s, just like Peter Turchin. And like Turchin Gavrilets is all over the place trying to produce formal models to elucidate a range of diverse questions. Gavrilets' research interests are social and cultural evolution, speciation and adaptive radiation, sexual conflict, holey fitness landscapes (this is how I knew him originally) and microevolutionary processes and macroevolutionary patterns. Good luck with all that, as they say....

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Punctuation Error?   posted by DavidB @ 4:59 AM
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Readers who lived through the Punctuated Equilibrium controversy of the 70s and 80s will recall that it petered out rather inconclusively, largely for lack of decisive empirical evidence one way or the other. The fossil record is seldom good enough to distinguish unambiguously between punctuational and gradual modes of evolution, one problem (noted already by Darwin) being that the sudden appearance of a new form in a given locality may result from migration rather than rapid evolution in the same place.

Given these difficulties, a disproportionate amount of attention was focused on a handful of examples that seemed to show good evidence either of punctuational or gradual evolution. One of the best examples on the punctuationist side of the debate was a study of molluscs in the Turkana Basin of Africa by P. G. Williamson [Note 1] Williamson's study was criticised at the time on various grounds - for example that the changes observed might be due to environmental stress rather than genetic evolution - but the critics did not produce new evidence from the field.

That is changed by an article [Note 2] by a Dutch team in a recent issue of the journal Evolution....



The Abstract of the article is as follows:

A running controversy in evolutionary thought was Eldredge and Gould's punctuated equilibrium model, which proposes long periods of morphological stasis interspersed with rapid bursts of dramatic evolutionary change. One of the earliest and most iconic pieces of research in support of punctuated equilibrium is the work of Williamson on the Plio-Pleistocene molluscs of the Turkana Basin. Williamson claimed to have found firm evidence for three episodes of rapid evolutionary change separated by long periods of stasis in a high-resolution sequence. Most of the discussions following this report centered on the topics of (eco)phenotypy versus genotypy and the possible presence of preservational and temporal artifacts. The debate proved inconclusive, leaving Williamson's reports as one of the empirical foundations of the paradigm of punctuated equilibrium. Here we conclusively show Williamson's original interpretations to be highly flawed. The supposed rapid bursts of punctuated evolutionary change represent artifacts resulting from the invasion of extrabasinal faunal elements in the Turkana palaeolakes during wet phases well known from elsewhere in Africa.


I have read the full article (available here), which looks convincing on this particular case (but what do I know about old African molluscs?) [Added: a more easily readable pdf version is also available. Google 'bocxlaer turkana' and you should find it.] The strongest point is that it is not just armchair criticism but based on extensive new fossil collecting. But since I specialise in armchair criticism I can hardly throw any stones.

Obviously one such case doesn't disprove punctuated equilibrium, but Williamson's study was in some ways the 'poster child' for the theory (more so than even Eldredge and Gould's own studies), so its demolition (if accepted) would be a serious blow.

Note 1: P. G. Williamson, 'Palaeontological documentation of speciation in Cenozoic molluscs from Turkana Basin', Nature, 1981, 293, pp.437-43. Also reprinted in Evolution Now, ed. John Maynard Smith, 1982. I can't find any publications by Williamson after 1990, and I believe I have read somewhere that he died at a sadly early age. My apologies if I am mistaken.

Note 2: Bert van Bocxlaer, Dirk van Damme, and Craig S. Feibel, 'Gradual versus punctuated equilibrium evolution in the Turkana Basin molluscs: evolutionary events or biological invasions?', Evolution, 2008, 62, pp.511-20.



The Mongol Art of War   posted by Razib @ 1:23 AM
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If Genghis Khan and the Making of the Modern World grated on you because of the transparent lack of scholarly objectivity, I recommend Timothy May's The Mongol Art of War. May usually attempts to present "both sides" in any given scholarly debate, but he also tells you which side is the majority and which the minority. And there's good quantitative data, like the fact that Mongol light cavalry had a range of up to 300 meters in terms of their bows. The Mongol Art of War makes it pretty obvious that courage is sometimes overrated as an ingredient of conquest, the Mongols rarely engaged in pitched battles because they weren't exceptional hand-to-hand fighters. Rather, when battling an enemy on open field they simply barraged their opponents with missile fire until attrition wore them down. Their reputedly high accuracy from long distances meant that they could stay out of danger while simultaneously inflicting casualties on the opposition. Not to be trite but it sounds like a precursor to "shock & awe" via air power medieval style.

It seems understandable that chivalry might emerge in societies where martial elites have incentives to formalize & codify and so minimize the risks inherent in the art of war, which is after all their primary profession. In contrast, the Mongol war machine which emerged in the early 13th century was notable for its relatively exceptional social egalitarianism. The Mongol army did not consist of an elite professional war-band, but rather was drawn from vast swaths of the adult male tribal population of Mongolia (on the order of perhaps 1/2 of the adult males served in the mobile armies during the initial years). Like the Roman legions before 100 BCE this was a nation of soldiers on the march, not the soldiers of a nation. Genghis Khan's light cavalry simply leveraged the typical skills of a nomad on a horse with bow in hand. The rapid expansion from the Yellow to the Black seas was due less to the calculated glory seeking of status seeking aristocrats than the random-walk rapaciousness of nomads whose lives had been characterized by existence on the margins of subsistence supplemented by raiding of surplus producing sedentary farmers. To some extent the emergence of the Mongol Empire was a series of raids writ-large.

Addendum: One thing I found interesting was the suggestion that one of the major reasons that Mongol expansion into the Middle East ran out of steam was lack of pasture for their horses. Each Mongol warrior might have had 5-15 horses. In South China the Mongols under Kublai Khan had to reinvent themselves because light cavalry did not offer any comparative advantage in the local ecology. And later Mongol attempts to expand into Southeast and Maritime Asia generally failed more often than not. In many parts of Eurasia the Mongols were defeated, but like the Romans before them they kept coming and eventually overcame resistance. This makes me wonder about true historical significance of the Mamluk defeat of the Mongols at the Battle of Ain Jalut, as in many accounts this is a great historical turning point. The implication is that if the Mongols had not been defeated in this battle they would have gone on to conquer all of North Africa. But as I alluded to above in Russia there were defeats but the Mongols bounced back. In contrast they were defeated several times by the Mamluks after Ain Jalut. This to me points to ecological constraints on the comparative advantage of the Mongol-way-of-war. Of course it is also quite plausible that empires have natural limits to their size contingent upon the scalability of communication lines as well as the diminishing returns on additional increments of territory.

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Monday, September 29, 2008

Graphs on the rise of scientific approaches to humanity   posted by agnostic @ 2:16 AM
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Well, with the first post and a response to criticisms out of the way, I'll conclude with the graphs on some ideas that are gaining in popularity in the study of mankind. Where it says "social sciences," I've only searched JSTOR for the following journal categories: anthropology, economics, education, political science, psychology, and sociology. The social sciences, basically. (And I've used appropriate neutral comparisons as before.) The reason is that if "heritability" increases in usage, that could be due to its use in genetics -- I want to see how popular it is when talking about humans. (As before, graphs have simple titles, while the full search terms are listed in an Appendix.)










Contrary to what you might think, since about 1950 academics have become increasingly interested in the genetic influence on human nature, reversing a period of decline from roughly 1930 to 1950. There is also an apparent cyclical pattern on top of the increasing trend. Just make sure you refer to the heritability of "cognitive ability" rather than of "IQ" (see below).

I've broken up the graphs on Darwin in the social sciences to make the trends clearer. There is an early phase in Victorian times when Darwin's thoughts were everywhere, especially in discussing human beings. Around the turn of the century, his ideas become less popular, as mentioned above. Around 1940, when his ideas come back due to the modern synthesis in biology, they become more popular in the social sciences as well. Indeed, since the mid-1940s, his ideas have only become more important to social scientists -- whether they like it or not.

Notice that while "IQ" goes through cycles about an increasing trend, its synonym "cognitive ability" shows exponential increase. I assume that this is because "cognitive ability" is not a politicized term, while "IQ" is, resulting in outbreaks of hysteria where many more people of any ideological background begin talking a lot about it.

The same is true of "sociobiology," which Leftist academics such as the Sociobiology Study Group tainted with negative political associations, compared to its synonym "evolutionary psychology." Now, someone will say that evolutionary psychology is different -- that it studies the mental, psychological processes rather than just observed behavior. But that's nonsense -- if you've read one of the many evolutionary psychology articles about digit ratios, waist-to-hip ratios, whether the female orgasm is adaptive, and so on, you know that mental processes and cognitive science models rarely come up, except in the study of vision.

Indeed, "evolutionary psychology" increases at just the time when "sociobiology" decreases, in the mid-1980s, showing that the former is simply replacing the latter as the preferred term.

As further evidence that a decline in usage means a decline in popularity, "evolutionary psychology" gets lots of hits in the 1890s when pioneers of psychology like William James were obsessed with integrating evolution and the study of the human mind, and takes a nosedive and lies dead once behaviorism takes over in psychology around the 1920s.

Because "evolutionary psychology" and "cognitive ability" are safe terms politically, these are the obvious choices for people who don't want to have water poured over their head at a conference -- and the data show this rational choice. Interest has continued to skyrocket, although people use different codewords. Nothing like this turned up in the first post because it is not political suicide to talk about postmodernism or Marxism in academia -- but just try bringing up "IQ". It is fascinating that academics can adhere to the ideas of Marx, Lenin, Trotsky, or Stalin and be taken seriously, while anyone who would do so for the ideas of Mussolini or Hitler would be made a total pariah. I wouldn't take either numbskull seriously, but most educated people will, perhaps grudgingly, give a free pass to those who revere the ideological or political figures associated with The Other Great Dictatorships and Mass Murders.

I've already made general observations in the first post, and they carry over here, especially the fact that the history of ideas seems so unaffected by the history of the entire outside world -- one more idea that Marx got wrong. There is clearly change, struggle between groups, and so on, but they are largely internal to academia. The future -- or the near-future anyway -- looks pretty bright for those interested in the biological approach to studying humans and their ways, and who believe things like IQ are important. Any students who are still considering the social constructionist, Marxist, feminist, or Whateverist approach should at least learn the new theories, if for no other reason than to be employable in 5 to 10 years. Hell, you might even consider it a kind of Pascal's Wager.

APPENDIX

Here are the search terms I used, once again searching the full text of articles and reviews:

"cognitive ability" OR "cognitive abilities"

"darwin*" NOT "social darwinism" NOT "social darwinist" NOT "social darwinists"

"evolutionary psychology" OR "evolutionary psychologist" OR "evolutionary psychologists"

"heritability" OR "heritable"

"IQ"

"sociobiolog*"

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Response to criticism on the death of academic -isms   posted by agnostic @ 1:57 AM
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My first post detailed the demise of wooly-headed theories in academia. In this post, I'll also address some common criticisms that have come up so far. In the third post, just above this one, I will look at a rival class of theories, namely the scientific and in particular biological approaches to studying humanity. The take-home message is that, while the Blank Slate theories are slowly being driven out of academia, new ones based on the biological sciences are becoming ever more popular. But let's start with the criticisms:

1) You're confusing popularity with accuracy, truth, etc.

I never said anything to this effect. I am just interested in whether certain theories are becoming more or less prevalent. Now, I happen to believe that in the case of, say, psychoanalysis or Marxism, the theories are becoming less popular because people realize that they're not very insightful. And certainly what I think is a great theory could become unfashionable for whatever reason. Whether you're celebrating or mourning the death of some theory, I don't care -- I just want to show whether it is or is not dying.

2) You didn't account for the lag between when an article is published and when it is archived in JSTOR.

I did do that, but I was only explicit about it in the comments to the first post. Journals in JSTOR have a "moving wall" between original and archived dates, with most having a lag of 3 to 5 years. Here is the distribution of lag times. By excluding data from 2003 onward, I've taken care of 88% of journals. And I don't want to hear a non-quantitative objection that "the remainder could be affecting the results" -- tell me what you think the data-point should be for, say 2001, and then derive how large of an effect the 12% of journals would have to have in order to get that value. We'll see how reasonable that sounds. Moreover, since no moving wall is greater than 10 years, any decline that started before 1998 is not subject to even this vague objection -- for example, Marxism, feminism, and psychoanalysis.

3) You don't have a neutral control case to show that Marxism is "really" decreasing in popularity.

I did admit in the first post that ideally we'd have the total number of articles that JSTOR has for a given year, and that we'd divide the number of articles with Marxism by the total number of articles to get a frequency or prevalence. We can estimate the total by searching for articles with some highly frequent word, such as "the", so that the number returned is very close to the total. For "the", this approach is almost guaranteed to work, since almost no article would slip through the net.

However, JSTOR has a list of highly frequent words that it doesn't allow. Still, not all common words are blocked. I consulted a frequency list compiled by Oxford Online, and chose the highest-ranking words there which are not blocked by JSTOR, though I excluded the personal pronouns and "people," since I don't expect those to show up much in hard science or social science journals. This gives the variants of "time," "know," "good," and "look."

So, I've estimated the total number of articles for a year by searching for "time" OR "know*" OR "good*" OR "look*", where the asterisk means the word-ending can vary. How closely this estimates the true total is not of interest -- the point is that it serves as a common, neutral yardstick to measure the change from one year to the next.

Interestingly, using this control has almost no effect on the shape of the graphs from the first post. That is because the increase in the total number of articles increases only linearly from about 1940 onward, whereas the articles on postmodernism increase or decrease exponentially -- and an exponential divided by a linear is still growing or dying very fast. I've redrawn the original graphs and posted them here because it's easier for me; sometime soon, I'll substitute them into the first post for the record.

The only change I make to my original observations is that social constructionism is not so obviously declining anymore, although it is plateauing and apparently declining since 1998. If I had to guess about its behavior after 2002, I would say it's downward simply because none of the other theories plateaued for very long -- they quickly hit a peak and declined, so a steady high value does not appear to be stable for such theories.

4) You're mistaking a decline in usage with a decline in belief -- once the idea becomes taken for granted, practitioners stop referring to it explicitly.

Just on an intuitive level, we know this is horseshit -- do physicists not use the words "gravity" or "electricity" anymore, or no longer refer to Newton? This objection exemplifies the problem with the average arts and humanities major: he is content to build a logically coherent argument without doing a quick reality check for its explanatory plausibility. I guess that's why they end up in law firms.

But to provide evidence that usage tracks belief, here are some graphs for hard science keywords. In the case of Darwin, I excluded articles on "social Darwinism," which appears to be a, er, social construction in academia. See here. I have data on academic scarewords like "biological determinism," and perhaps in a future post I'll show those. Right now, I want to focus on articles that are at least somewhat level-headed. For ease of inspection, I've given each graph a simple title, and list the search terms at the end of this post in an Appendix.






As the disciplines of population genetics and sociobiology have become staples of biology, mentioning them by name has not declined -- just the opposite. Because they are such thriving fields, writing about them explicitly has shot up. Darwin's thoughts were immensely popular in Victorian times, but they languished because no one could tell how to unite them with the study of heredity. That was, until the modern evolutionary synthesis, which began in the late 1930s -- since then, interest has exploded. The same goes for Mendel's thoughts -- no one knew what the physical basis for his "gene" idea was, until the relationship between the genetic code and DNA was laid out in the late 1950s.

This shows that even hard science ideas can rise and fall and rise again, in these cases probably because some key aspect was found unsatisfying, until a later discovery fixed the problem, allowing the idea to become popular again. So there's hope for the unemployed psychoanalyst yet, assuming he can stick around for a half-century.

APPENDIX

Here are the search terms I used:

"population genetic" OR "population genetics" OR "genetics of populations"

"sociobiolog*"

"darwin*" NOT "social darwinism" NOT "social darwinist" NOT "social darwinists"

"mendel" OR "mendelian*" OR "mendelis*" NOT "mendelss*"

I put the last restriction on the Mendel search because I got a lot of results about the composer Felix Mendelssohn.

As with the first post, I searched the full text for both articles and reviews.

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Sunday, September 28, 2008

Feeling sleepy?   posted by p-ter @ 5:06 PM
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Nature Genetics this week has published a genome-wide association study of narcolepsy in the Japanese population. The finding in the paper is a variant that confers a modest risk of narcolepsy, but personally, I was blown away by Figure 1, reproduced above. The figure shows the strength of association of each of 500,000 SNPs with narcolespy, and the novel reproducible finding is on chromosome 22 (ie, it doesn't stand out all that impressively in this plot). The major signal, absolutely swamping everything else, is in fact in the MHC region (called HLA in humans).

This region, of course, contains risk factors for type I diabetes, crohn's disease, and most (all?) other autoimmune diseases. A quick google confirms that, indeed, thinking of narcolepsy as an autoimmune disease is not new, but it's definitely new to me, and it's pretty striking to see just how much more important the risk factors in HLA are compared to everything else.

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Saturday, September 27, 2008

R. A. Fisher and the Adaptive Landscape   posted by DavidB @ 6:01 AM
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In my note on Sewall Wright's concept of the Adaptive Landscape I said that I would later discuss R. A Fisher's views on the subject. Some commentators have claimed that Fisher held a definite view on the 'shape' of the landscape. For example, a book by Sergey Gavrilets includes a section on 'Fisher's single-peak fitness landscapes', with the claim that:

In contrast to Wright, Fisher... suggested that as the number of dimensions in a fitness landscape increases, local peaks in lower dimensions will tend to become saddle points in higher dimensions. In this case, according to Fisher, natural selection will be able to move the population without the need for genetic drift or other factors. A typical fitness landscape implied by Fisher's views has a single peak. - Gavrilets, p.36


I think this goes beyond anything that Fisher actually says about Wright's adaptive landscape. There is of course room for debate about what an author's views imply. My own interpretation is that Fisher was sceptical about the value of the landscape concept as such, because both environmental and genetic conditions were too changeable for the metaphor of a 'landscape' to be useful. For Fisher the question of the 'shape' of the landscape therefore did not arise as a major issue, and he had no need to take a firm view on it. I discuss this interpretation below the fold.


Sources

As I pointed out in my earlier note, Wright himself seldom if ever used the term 'landscape', so we should not expect to find the term in Fisher either. Wright usually referred to a 'field' of gene combinations, and a 'surface' of selective values. He used these concepts mainly to illustrate his shifting balance theory of evolution. Any comments by Fisher that are relevant to the shifting balance theory could therefore also be relevant to the landscape concept. Even with this broad scope, I can find few published comments by Fisher on the subject. The main ones are in his 1932 review of Wright's paper on 'Evolution in Mendelian Populations', reprinted in Bennett (ed.), his 1941 paper on 'Average excess and average effect of a gene substitution', his 1953 paper on 'Population genetics', and his 1958 paper on 'Polymorphism and natural selection', all available at the Fisher Archives here.

In addition to Fisher's published writings, his correspondence contains a few relevant remarks. Most of his correspondence is accessible at the Fisher Archives, and a good selection of his letters on evolution and genetics is published in Bennett (ed.) Two letters are especially relevant. In February 1931 Wright outlined his landscape concept in a letter to Fisher, quoted in Provine's biography of Wright (p.272). In a reply Fisher made some sceptical comments. Then in 1938 Fisher's colleague E. B. Ford described Wright's concept in a popular book on genetics. In a letter of 2 May 1938 to Ford, commenting on his book, Fisher gave what is probably his longest critique of the landscape concept. The letter is published in Bennett (ed.) (p.201-2) and available at the Fisher Archives, so I will not quote it in full, but it should certainly be read by anyone interested in this issue.

From Fisher's published and unpublished writings we can extract a number of criticisms of Wright's theory.

The interpretation of the dimensions of the landscape

In his biography of Wright, William B. Provine has pointed out that Wright in various places used two different interpretations of the genetic 'dimensions' of the landscape, which in Provine's view are inconsistent (Provine, p.313). In one interpretation the dimensions represent the number of alleles of a given type in an individual genome, while in the other interpretation they represent the frequency of those alleles in a population. Provine points out that in the first interpretation there is properly speaking no continuous surface, but only a lattice of discrete points. He also argues that there is no way of validly transferring conclusions from one interpretation to the other. I believe that these criticisms are somewhat overstated, but it is interesting to find that they are both anticipated by Fisher. In his letter to Ford, Fisher comments that either Ford's description of Wright's views, or the views themselves, are confused, and points out that 'so far as individuals are concerned, there is only a discontinuous aggregate of lattice points, each having its own selective value. There is no continuum of possible values in which we might speak of peaks or maxima.' In his article of 1941, Fisher also criticises one of Wright's own accounts, remarking that Wright 'confuses the number of genotypes, e.g. 3^1000, which may be distinguished among individuals, with the continuous field of variation of gene frequencies.... the large number of genotypes gives no reason for thinking that even one peak, maximal for variations of all gene ratios should occur in this field of variation' (1941, p.378). It is surprising that no-one else seems to have picked up on the apparent confusion in Wright's accounts until Provine's book in 1986.

The number of peaks

As discussed in my earlier post, Wright believed that there are usually a very large number of local fitness maxima in the landscape. Fisher, on the other hand, believed that this was unproven. As noted above, he thought that Wright's view was partly due to confusion between optimal genotypes and optimal frequencies. There is no easy transition from the existence of multiple optima among genotypes to multiple optima among frequencies. I have suggested in my earlier post that in some circumstances (notably where the optimal genotype is homozygous at all loci, and fitness is not frequency-dependent) there can be such a transition, but this is a special case. In general Fisher was correct to regard Wright's argument as inconclusive.

Fisher makes another criticism in his letters to Wright and Ford. In the letter to Wright he says:

In one dimension a curve gives a series of alternative maxima and minima, but in two dimensions two inequalities must be satisfied for a true maximum, and I suppose that only about one fourth of the stationary points will satisfy both. Roughly I would guess that with n factors only 2^-n of the stationary points would be stable for all types of displacement, and any new mutation will have a half chance of destroying the stability. This suggests that true stability in the case of many interacting genes may be of rare occurrence, though its consequence when it does occur is especially interesting and important.


In his letter to Ford, Fisher writes:

In one dimension, as in a road, we pass over an alternative series of hills and dips, so that half of the level points are maxima. In two dimensions, in addition to peaks and bottoms we have cols [i.e. saddle points], which may be regarded as the lowest points on ridges or the highest points on valleys, the curvature of the ground being positive in one direction and negative in another, and the peaks are only about a quarter of the level spots. In n dimensions only about one in 2^n can be expected to be surrounded by lower ground in all directions.


Disregarding for a moment the important comment in the first letter about new mutations, Fisher's thinking seems to be as follows. In each dimension of gene frequencies, only about half of the level points will be maxima. Assuming that the location of the maxima in each dimension is independent of the other dimensions, the probability that a level point will be simultaneously maximal in all dimensions will only be about (1/2)^n, or 1 in 2^n.

As these are just comments in private letters, it is difficult to know how much weight we should put on them. Fisher uses the words 'roughly', 'guess', and 'about', which do not suggest a dogmatic position. The validity of the two key assumptions - that about half of the level points in each dimension will be maxima, and that these will be independent of each other - could be discussed at length. But even at best, Fisher's argument only goes to show that the proportion of the level points which are all-round maxima will fall as the number of dimensions increases (which, incidentally, Wright himself accepted, e.g. at ESP p.226). It does not follow that the number of all-round maxima will remain small. If Fisher believed that this was necessarily the case (which is not clear), he was mistaken. It is quite possible that with an increasing number of dimensions the number of level points may increase faster than the proportion of all-round maxima declines. Indeed, it has been claimed that this is generally the case, but this is also unproven. (I will discuss this more fully in a separate post.)

I have not found any definite statement by Fisher either accepting or denying the existence of multiple optima. As I pointed out in my post on Fisher's views on epistasis, he accepted that there could be alternative stable allele frequencies at particular loci. As far as I can see, Fisher would not have denied in principle the possibility of multiple optima for the genome as a whole, and indeed his 1931 letter to Wright might be interpreted as accepting them as an important if rare phenomenon. But overall I think Fisher's position should be described as deeply sceptical. Wright himself said that Fisher 'did not accept the concept of multiple selective peaks' (Wright,1970, p.23), which is literally true, provided it is not taken as implying outright rejection either.

The mean fitness of the population

In Wright's theory, a population is expected to 'climb' up the slope of the fitness landscape under the influence of natural selection, implying that the mean fitness of the population increases. (Selection may however be offset by migration, recurrent mutation, or genetic drift.) In his publications from 1935 onwards (e.g. ESP p.239, 366) Wright uses a formula which may be expressed as delta-q = [q(1 - q)/2W][dW/dq], where q and (1 - q) are the frequencies of two alleles, delta-q is the single-generation change in q, W is the mean fitness of the population, and dW/dq is the partial derivative of W with respect to changes in q. The formula may be interpreted as saying that the effect of selection on the frequency of a particular allele is proportional to its effect on the mean fitness of the population (as well as to the current frequency distribution q(1 - q)).

In his 1941 paper Fisher strongly criticised this formulation, showing by a somewhat roundabout argument that it depends on the assumption of random mating, and claiming that any attempt to relate selection pressure to mean fitness is 'foredoomed to failure just so soon as the simplifying, but unrealistic, assumption of random mating is abandoned' (p.378). Wright's derivation of his formula, e.g. at ESP p.239, does indeed assume random mating. But Fisher's objection is not just technical: 'In regard to selection theory, objection should be taken to Wright's equation principally because it represents natural selection, which in reality acts upon individuals, as though it were governed by the average condition of the species or inter-breeding group. Early selectionists, following in this respect the language of the earlier theological writers on organic adaptation, often speak of selection as directed 'for the good of the species'. In reality it is always directed to the good, as measured by descendants, of the individual. Unless individual advantage can be shown, natural selection offers no explanation of structures or instincts which appear to be beneficial to the species. Yet in Wright's equation the whole evolutionary sequence would appear to be governed by the principle of increasing the 'general good'.' (p.378) I think this is somewhat unfair to Wright, who did not ascribe any causal efficacy to the fitness of the population as such, but Fisher's statement is important as his first general criticism of 'good of the species' thinking. He makes similar criticisms in his 1953 and 1958 papers. In the 1958 edition of GTNS a section on 'The Benefit of the Species' is added, which has become highly influential on modern evolutionary thinking. Although this new section does not refer to Wright, it is plausible that Fisher's sharpening of his hostility to 'good of the species' thinking was stimulated by his objections to Wright's equation.

New Mutations

As already mentioned, in his 1931 letter to Wright, Fisher argues that 'any new mutation will have a half chance of destroying the stability' of an optimal gene frequency. He makes a similar point in his published review of Wright's 1931 paper on 'Evolution in Mendelian Populations', saying that 'even under static conditions, unless it is postulated that the organism is as well adapted as it could possibly be (in which case, obviously, evolutionary improvement is impossible), the equilibrium will be broken by the occurrence of any favourable mutation, of which a steady stream will doubtless occur in one or other of the very numerous individuals produced in each generation. The advantage of the large populations in picking up mutations of excessively low mutation rate seems to be overlooked [by Wright]'.

Their attitude towards new mutations is one of the fundamental dividing lines between Wright and Fisher. Wright repeatedly played down the importance of favourable new mutations, on the grounds that their chance of occurring would be negligible even over long periods (see e.g. ESP pp.150, 165, and 321). He seems to have believed that all possible mutations would already have occurred often enough to be selected if they were favourable, so that the possibility of improvement through new mutations would already have been exhausted. Fisher, in contrast, believed that in large populations even very low mutation rates (say, of one in a thousand million per generation) could not be neglected, and that on an evolutionary time-scale of hundreds or thousands of generations they would provide scope for continuing evolution. It may of course be thought that neither Wright nor Fisher, in the 1930s, knew enough about the nature of genes to have any good basis for their opinions.

Changing Environment

Wright's concept of the adaptive landscape is explicitly based on the assumption of constant environmental conditions. Any change in those conditions involves a change in the landscape itself. Wright was of course aware that environments could change, but he seems to have regarded the 'landscape' as having an underlying continuity of existence even if environmental fluctuations might temporarily change its shape. (I will consider Wright's views on this further in my final post on the shifting balance theory.)

Fisher, on the other hand, believed that environmental change was in one sense irreversible. In the section 'Deterioration of the Environment' in GTNS he emphasised especially the organic environment of competitors, etc:

For the majority of organisms... the physical environment may be regarded as constantly deteriorating... Probably more important than the changes in climate will be the evolutionary changes in progress in associated organisms. As each organism increases in fitness, so will its enemies and competitors increase in fitness; and this will have the same effect, perhaps in a much more important degree, in impairing the environment, from the point of view of each organism concerned. - The Genetical Theory of Natural Selection, Variorum Edition, ed. Henry Bennett, 1999 p.41-2


In his review of Wright's 'Evolution in Mendelian Populations' (reprinted in Bennett, ed.) Fisher again emphasised environmental change:

Professor Wright considers that: 'In too large a freely interbreeding population there is great variability, but such a close approximation to complete equilibrium of all gene frequencies that there is no evolution under static conditions'. He therefore argues that the subdivision of species into partially isolated local races of small size is an important condition not merely, as is obvious, for fission into distinct species, but for progressive evolution. This conclusion is much more debatable [Fisher then makes his point about the importance of new mutations even under static conditions]... Moreover, static conditions in the evolutionary sense certainly do not occur, for, apart from geological and climatological changes, the evolutionary progress of associated organisms ensures that the organic environment shall be continually changing


In short, as several recent commentators have noted, Fisher held a 'Red Queen' conception of evolution, in which organisms have to keep constantly running just to keep up with the competition. This is quite alien to Wright's conception, in which under the influence of selection alone the organic world would soon grind to an evolutionary halt. The extent to which either of these views is correct is a matter for empirical observation. Genetic studies of living populations tend to show continual change, at least at a microevolutionary level, which might seem to support Fisher's view, whereas paleontologists often claim to observe long-term stasis in morphological traits, which might support Wright. This is of course one of the points at issue in the debate over 'punctuated equilibrium', which seems to have petered out through boredom (and the death of some key participants) rather than being resolved. A possible explanation of the apparent conflict of evidence is that traits in hard body parts may be more tightly constrained by stabilising selection than biochemical and behavioural traits. For other suggestions see Williams, Chapter 9.

Refs:
J. H. Bennett, ed.: Natural Selection, Heredity and Eugenics: Including selected correspondence of R. A. Fisher with Leonard Darwin and others, 1983.
Sergey Gavrilets, Fitness Landscapes and the Origin of Species, 2004.
William B. Provine, Sewall Wright and Evolutionary Biology, 1986.
Sewall Wright: Evolution: Selected Papers (ESP), ed. William B.Provine, 1986.
George C. Williams: Natural Selection: Domains, Levels, and Challenges, 1992.
Sewall Wright: 'Random drift and the shifting balance theory of evolution', in Mathematical Topics in Population Genetics, ed. Kojima, 1970.


Friday, September 26, 2008

10 Questions for Parag Khanna   posted by Razib @ 11:59 AM
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Parag Khanna is the author of The Second World: Empires and Influence in the New Global Order. He is also Director of the Global Governance Initiative and Senior Research Fellow in the American Strategy Program at the New America Foundation. His website is Paragkhanna.com, where one can find a repository of articles, videos and interviews. Below are 10 questions. (in case readers are curious, I did read The Second World in one sitting)


1) Another recent work which I think one can compare to your book, "The Second World: Empires and Influence in the New Global Order," is Fareed Zakaria's "The Post-American World." If I had to contrast the two I would suggest that Fareed's narrative is both broader in scope and thinner in detail. "The Post-American World" attempts to describe a possible future trajectory for the whole world while you focus specifically on the Second World (albeit, a rather a large canvas in and of itself). And while Fareed tends to utilize simple and general frameworks (e.g., China and India do not believe in God), you seem to rely on more thick description empirically (balancing both quantitative statistical data with on-the-ground observation) as well as a more scholarly theoretical superstructure (such as H. L. Mackinder's "Heartland" model). Would you say this is a fair description of the differences?

*** My "Second World" is certainly broader in scope than "Post-American World" in that it covers literally the entire planet (but with some areas like Central America and Sub-Saharan Africa getting very short treatment). He gives more space to India than I do, and we both accord much attention to China, while I add in China's role in key regions like Latin America, the Middle East, and Central Asis as well. Even though my book is a travelogue with micro-detail, Fareed and I have nearly identical conclusions about the growing confidence of "The East" and the Second World more broadly (which he calls "the rest"). I wanted to have the scholarly superstructure in my book because in addition to providing descriptive detail, I wanted to be predictive about what the rise of new powers will do to geopolitical transition, the balance of power, the future constraints on American foreign policy, and so on.


2) You make copious reference to geographer H. L. Mackinder and those who responded to his hypothesis about the centrality of the Eurasian core in world domination (e.g., Nicholas J. Spykman and the "Rimland"). As a self-identified geography-nerd I can say that your references to grand theoretical frameworks were deftly integrated into the narrative. On the other hand, can you expand on the value which these sorts of models may give to the typical lay reader? Specifically, do you believe that the theory allows one to plausibly stitch together the copious data which you present within your narrative a more comprehensible manner?

*** Even in the age of technology and globalization, geography is still destiny for most. I try to demonstrate just how important the regional context is for evaluating a country's situation and options -- it is more important than the global in most cases. So indeed, Mackinder and geopolitics' emphasis on population, resources, location, sea access, natural barriers, and other features of geography remain absolutely pivotal to understand a country's prospects. Let's take the biggest debate in Asia today (from America's point of view), namely India vs. China. Just because they both have over 1 billion people, that does not make them equal. Even if they both had efficient regimes (which China does and India doesn't), or even if the regimes were reversed, and India had the "better" government, India would still face the reality that it is hemmed in by the Himalayan mountains (the world's tallest) and vast oceans, and has acrimonious relations (at best) with all its neighbors. This severely limits its power projection capability. China, on the other hand, borders more countries than any other in the world, which is extremely useful when spreading influence by either economic, demographic, military, or infrastructural means.


3) I recently read "After Tamerlane: The Global History of Empire Since 1405," which seems to argue there was a fundamental shift in the rise and fall of polities around this period. Specifically, the author seems to be making the case that the Gunpowder Empires fundamentally reversed the power dynamic which had long privileged the peoples of Mackinder's Heartland over the settled societies of Spykman's "Rimland." The defeat of the Dzunghar Confederacy by the Manchus and the rollback of the Tatar by the Russian Empire come to mind. Since I am not fluent in Mackinder's ideas at anything more than a caricature level, am I right to believe that he his argument was one of strategic control of territory, as opposed to the dynamic forces of history being driven by peoples shaped by the ecology of the Heartland itself?

*** "After Tamerlane" is a wonderful work of scholarship. Both the factors you identify -- strategic control of territory and the ecology of the Heartland -- were important for Mackinder. It wasn't just that the Heartland would be impervious to naval attack/control, but also that it possessed rich natural resources (water, timber, etc.) In fact, based on differing understandings of Mackinder's emphasis, scholars have come up with different geographies for the precise location of "Heartland" and "Pivot", Mackinder's other key geographic concept.


4) Reading "The Second World" I felt the shadow of books such as "Guns, Germs and Steel" and "The Wealth and Poverty of Nations." In other words, fixed geographical parameters interact dynamically with historical contingencies to shape the patterns of variation we see around us. For example, geography does not mean that Argentina is a wealthy land, but, if history is a guide it suggests that it can be a wealthy land because of the potential productivity of agriculture in its particular climate. I believe these coarse marco-level parameters are critical and do add value in our attempt to model the reasons for the shape of the past, the nature of the present, and the possible trajectories of the future. But I also have an interest in biology, and I am of the opinion that economists for example would gain value by deviating from the uniform Homo economicus assumption and take into account individual and group differences. There are strong indications for example that Toxoplasma gondii, a parasite which one may catch from cats, can change personality and predisposition and generate between-cultural differences. There is also data which suggests that personality variation may be controlled by genes which modulate dopamine pathways. Finally, there are also new avenues of research suggesting genetic variation which controls differences between individuals in behavioral economics experiments. These are simply three examples. I believe in the importance of geography as a major macroscale parameter, but, I am also one who suspects that many of these coarse differences we see across the world may be rooted in microscale variation. Some economists are moving into this domain and attempting to find causal connections between the micro and macroscale. Do you know if scholars in international relations community have taken notice? Or are there just too many other low hanging fruit to analyze so that it is impractical at this moment to integrate these domains into the field?

*** You might have noticed that I try to pitch the book as a work of "geopolitical psychology" and I use quite a few metaphors from sociology as well. I strongly avoid any rational actor models/biases in the book, and in fact try to highlight "irrational" behavior wherever possible and show it as a product of history/culture/geography: just take Chavez in Venezuela, Gaddafi in Libya, and Putin in Russia, three examples I delve into in the book. I also argue that a nation's psychology is schizophrenic (particularly in the Second World), and that nations can be satiated in a manner that tracks to Maslow's famous "hierarchy of need". But yes, beyond game theory there is increasing amounts of work in political science that looks at such non-social sciene approaches to undetstanding behavior in the political arena as well.


5) I was struck a bit by the Sinocentric focus of much of the book. China looms large. In contrast, you don't spend much time on India, asserting that it is basically a Third World nation, and will remain one for some time. This seems plausible to me skimming over the data on any human development index (or, the fact that it seems likely that the majority of the world's mentally retarded due to nutritional deficiency, cretins, reside on the Indian subcontinent). Nevertheless, the media in the United States has constructed a China vs. India narrative. Good copy? Or do some people actually think in these terms? (as I suggest above, it seems that the comparison is laughable looking at the bottom-line statistics on most vital indices)

*** In addition to my answer above about how geography impacts the China vs. India debate, it should be added, in agreement with you, that it makes good copy. After all, how else could one come up with an acronym like "Chindia" or "BRICS", both of which actually speak against the argument of rivalry. In any case, a 19th-century view of the balance of power would certainly suggest that India would make for a strong, populous, democratic, industrial, nuclear, naval superpower partner in the quest to contain China, and thus it's a very useful construction on the part of the U.S. Pentagon. That said, a great deal more depth has been added to the US-India relationship over the years, especially in the IT and now biotech and other areas, so there is a pattern of growing trust between the two since the end of the Cold War, and independent of the military relationship.


6) Your book was published last year. Events move fast. You spend some time on Georgia, and it is not a particularly flattering picture. A friend of mine told me several weeks ago the basic outline you present, which I would characterize by suggesting that Georgia lay somewhere in the great middle between Bangladesh and Finland in corruption and the robustness of civil society. But during the recent course of events I heard little detail of the nation of Georgia as opposed to the specific blow-by-blow of events (or what we know) involving South Ossetia, Russia and Georgia. Is simply an unchangeable feature of the media, or a bug which might be fixed in future releases? Is there any way we can prevent this? Geographical knowledge isn't a top priority now...but it seems that a little data would go a long way in making more informed foreign policy decisions.

*** It certainly would! And that is why my book attempts to be an inside-out look at Georgia. I present Mikhael Saakashvili as a corrupt, power-hungry and pugnacious semi-autocrat, and Georgia as a West African micro-state in the Caucasus. I talk about the poor roads, the belching buses, the squalid villages and the arrogant government. If more foreign policy experts and the general public understood these things earlier on, we would have heard less bullish talk about Georgia I think.


7) You lived in the United Arab Emirates at some point and profile Dubai. So quick question, is Dubai sustainable over the next decade? There are some questions about how over-leveraged and how it is being bankrolled by taking on debt. Additionally, as you allude to in "The Second World" it also extracts labor productivity rather cheaply out of most of its South Asian workforce and it seems there is a likelihood that at some point in the near future the cost of this labor might increase because of the imposition of what we in the States would term humane working conditions.

*** I do believe that Dubai is sustainable over the next decade - and the entire UAE even more so. The country has quickly taken up an essential place as a node in the globalized world, both financial (think SWFs), geographical (a key re-export zone located between Europe and Asia), and political (a neutral and safe place in a turbulent region). There is an outside view of labor conditions and an inside view. The outside view equates third world/Asian labor conditions as tantamount to slavery. The inside view shows that they're considering a proper minimum wage, are aquiring low-cost but energy-efficient housing in the labor camps, and that the workers are there because they want to be there and earn enoughto make the UAE/Gulf the second largest source of global remittances (behind the US). So there will be bumps in the road, but Dubai is the Arab world's first "global city" and absolutely essential for the region and now the world.


8) Speaking of working conditions and cheap labor, in the closing of the book you make reference to immigration in the United States. There are some, quite often economists, who make a case for the enriching value of open borders and free movement of labor, while others would like to close borders in the interests of cultural homogeneity and tightening labor supply to increase wages. Immigration has been a major flashpoint here in the United States over the past few years, and we haven't really resolved anything and seem to be tabling the issue for now. If you could design a system of immigration for the United States what would it look like?

**** The notion of closed borders vs. total free movement represent two extreme bookends, neither of which is realistic. The balance has to be found between bringing in sufficient low-cost Latin labor to do the work that Americans won't do, while also bringing the workers up the value chain so that real wages are pulled down. I do think bringing the currently illegal population "above board" as Schwarzeneggar proposed a while back is a good idea - it will help to get a better accounting of numbers of immigrants/workers in the country and legitimize their presence.


9) Empires loom large in the actions of Second World powers, the United States, China and the EU. I get the sense from the book that you think that the EU has acquitted itself rather well in the world of late in terms of advancing its own interests, both in terms of realpolitik and in the domain of spreading its normative outlook. Additionally, you observe that the EU allows for both unity and diversity; nations can preserve their language and culture upon admission, though obviously centralizing and homogenizing trends are also apparent. Of late some scholars have been looking back to empires of the past as models for diversity existing cheek-by-jowl with political unity. But I would assert that despite diversity most empires of the past were dominated by one identity. For example, Polybius famously observed that the power of the Roman state was its assimilative capacity, but Anastasius in the late 5th century was probably the first emperor who self-identify as a Hellene. Emperors of "exotic" lineage such as Philip the Arab or Septimius Severus (Punic on his father's side) were Latinized. In "The Fall of the Roman Empire: A New History of Rome and the Barbarians" the author asserts that the liberal education which was the norm among Roman aristocrats was essential in inculcating not just specific values, but an upper class Latin accent which could mark one's social origins through life immediately upon first contact. So my point is that these diverse empires had herrenvolk. You seem to point to China's Han ethnicity as something of this sort, but the Han are on the order of 90% of the population of their state. In contrast, there is no such preponderance of ethnicities in the EU. Could it be that past exemplars are simply not applicable to the present? In "The Great Upheaval: America and the Birth of the Modern World, 1788-1800" Jay Winik contends that it was a common assumption during the 18th century that popular governmental forms such as republics and democracies were simply not scalable beyond the city-state, and yet here we are over 200 years later.

*** In her excellent book "Day of Empire," Amy Chua explains how tolerance of diversity was a renewing force for major historical empires and becomes the lifeblood of sustainability, even as it eventually can bring down an empire. The interplay of technology and historical learning is what has allowed the EU to become the modern day Holy Roman Empire so successfully.


10) You offer that you've traveled to over 100 nations in "The Second World." Certainly impressive, but I'm curious as to the range of linguistic fluency necessary make yourself understood. Was English sufficient, or did you have to lean on languages which you'd learned or already knew besides English? Were there interregional differences?

*** One needs a different linguistic strategy for each region. In Eastern Europe I got by with a mix of German (which I speak fluently), English, and some translators for Ukrainian and Russian. I speak Spanish so was okay in South America. In the Mideast I used basic Arabic to get by on the street, but did interviews either in English or with translators. And in China I needed translators all the time. If I could have done it all over again it would have been nice to speak Russian given all time I spent in Central Asia, but I wouldn't recommend to young students today to learn Russian. I think it is best to learn Arabic or Chinese (or both) today.

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Surfwise on Netflix   posted by Razib @ 12:21 AM
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If you have Netflix, the Paskowitz family documentary Surfwise is now watchable online.