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ResearchBlogging.org One of the most persistent debates about the process of evolution is whether it exhibits directionality or inevitability. This is not limited to a biological context; Marxist thinkers long promoted a model of long-term social determinism whereby human groups progressed through a sequence of modes of production. Such an assumption is not limited to Marxists. William H. McNeill observes the trend toward greater complexity and robusticity of civilization in The Human Web, while Ray Huang documents the same on a smaller scale in China: A Macrohistory. A superficial familiarity with the dynastic cycles which recurred over the history of Imperial China immediately yields the observation that the interregnums between distinct Mandates of Heaven became progressively less chaotic and lengthy. But set against this larger trend are the small cycles of rise and fall and rise. Consider the complexity and economies of scale of the late Roman Empire, whose crash in material terms is copiously documented in The Fall of Rome: And the End of Civilization. It is arguable that it took nearly eight centuries for European civilization to match the vigor and sophistication of the Roman Empire after its collapse as a unitary entity in the 5th century (though some claim that Europeans did not match Roman civilization until the early modern period, after the Renaissance).

It is natural and unsurprising that the same sort of disputes which have plagued the scholarship of human history are also endemic to a historical science like evolutionary biology. Stephen Jay Gould famously asserted that evolutionary outcomes are highly contingent. Richard Dawkins disagrees. Here is a passage from The Ancestor’s Tale:

…I have long wondered whether the hectoring orthodoxy of contingency might have gone too far. My review of Gould’s Full House (reprinted in A Devil’s Chaplain) defended the popular notion of progress in evolution: not progress towards humanity – Darwin forend! – but progress in directions that are at least predictable enough to justify the word. As I shall argue in a moment, the cumulative build-up of compelx adaptations like eyes strongly suggest a version of progress – especially when coupled in imagination with of the wonderful products of convergent evolution.

Credit: Luke Jostins
Credit: Luke Jostins

One of those wonderful products is the large and complex brains of animals. Large brains are found in a disparate range of taxa. Among the vertebrates both mammals and birds have relatively large brains. Among the invertebrates the octopus, squid and cuttlefish are rather brainy. The figure to the right is from Luke Jostins, and illustrates the loess curve of best fit with a scatter plot of brain size by time for a large number of fossils. The data set is constrained to hominins, humans and their ancestors. As you can see there is a general trend toward increase cranial capacities across all the human populations. Neandertals famously were large-brained, but they exhibited the same secular increase in cranial capacity as African Homo. On the scale of Pleistocene Homo and their brains the idea of the supreme importance of contingency seems ludicrous. Some common factor was driving the encephalization of humans and their near relations over the past two million years. This strikes me as very strange, as the brain is metabolically expensive, and there are plenty of species with barely a brain which are highly successful. H. floresiensis may be a human instance of this truism.

But what about the larger macroevolutionary pattern? Is there a trend toward larger brain sizes in general, of which primates, and humans in particular, are just the most extreme manifestation? Some natural historians have argued that there is such a trend. But, there is a question as to whether increased brain size is simply a function of allometry, the pattern where different body parts and organs tend to correlate together in size, but also shift in ratio with scale. The nature of physics means that very large organisms have to be more robust because their mass increases far faster than their surface area. By taking the aggregate relationship between body size and brain size, and examining the species which deviate above or below the trend line, one can generate an encephalization quotient. Humans, for example, have a brain which is inordinately large for our body size.

And yet there are immediate problems looking at relationships between body and brain size, and inferring expectations. Different species and taxa are not interchangeable in very fundamental ways, and so a summary statistic or trend may obscure many fine-grained details. A new paper in PNAS focuses specifically on various mammalian taxa, corrects for phylogenetics, and also relates encephalization quotient by taxa to the proportion of social animals within each taxon. Encephalization is not a universal macroevolutionary phenomenon in mammals but is associated with sociality:

Evolutionary encephalization, or increasing brain size relative to body size, is assumed to be a general phenomenon in mammals. However, despite extensive evidence for variation in both absolute and relative brain size in extant species, there have been no explicit tests of patterns of brain size change over evolutionary time. Instead, allometric relationships between brain size and body size have been used as a proxy for evolutionary change, despite the validity of this approach being widely questioned. Here we relate brain size to appearance time for 511 fossil and extant mammalian species to test for temporal changes in relative brain size over time. We show that there is wide variation across groups in encephalization slopes across groups and that encephalization is not universal in mammals. We also find that temporal changes in brain size are not associated with allometric relationships between brain and body size. Furthermore, encephalization trends are associated with sociality in extant species. These findings test a major underlying assumption about the pattern and process of mammalian brain evolution and highlight the role sociality may play in driving the evolution of large brains.

A key point is that the authors introduce time as an independent variable, so they are assessing encephalization over the history of the taxon. This is clearly relevant for humans, but may be so for other mammalian lineages. The table and figures below show the encephalization slope generated by using time and body size as the predictors and brain size as the dependent variable. A positive slope means that brain size is increasing over time.

[nggallery id=21]

Two major points:

- Note that the slope is sensitive to the level of taxon one is examining. A closer focus tends to show more variance between taxa. So, for example, humans distort the value for primates in general. Bracketing out anthropoids paints a more extreme picture of encephalization, a higher slope. In contrast, the lemurs and their relatives exhibit less encephalization over time.

- The correlation between proportion of species which exhibit sociality and encephalization of the taxon is strong. From the text:

Encephalization slopes were correlated with both the proportion of species with stable groups (order R = 0.92, P = 0.005, n = 6; suborder R = 0.767, P = 0.008, n = 9; Fig. 2 A and B) and the proportion in either facultative or stable social groups (order R = 0.804, P = 0.027, n = 6; suborder R = 0.63, P = 0.04, n = 9).

The last figure makes it is clear that the correlations are high, so the specific values should not be surprising. Don’t believe these specific figures too much, how one arranges the data set or categorizes may have a large effect on the p-value. But the overall relationship seems robust.

266px-Alienigena
A highly encephalized “alien”

What to think of all of this? If you don’t know, one of the authors of the paper, Robin Dunbar, has been arguing for the prime importance of social structure in driving brain evolution among humans for nearly twenty years. The relationship is laid out in his book Grooming, Gossip, and the Evolution of Language. Robin Dunbar is also the originator of the eponymous Dunbar’s number, which argues that real human social groups bound together by interpersonal familiarity have an upper limit of 150-200. He argues that this number arises because of the computational limits of our “wetware,” our neocortex. Those limits presumably being a function of biophysical constraints.

One interesting fact though is that the median cranial capacity of our species seems to have peaked around one hundred thousand years ago. The average human today has a smaller brain than the average human alive during the Last Glacial Maximum! (see this old post from Panda’s Thumb, it’s evident in the charts) This may be simply due to smaller body sizes in general after the Ice Age. Or, it may be due to the possibility that social changes with the rise of agriculture required less brain power.

Ultimately if Dunbar and his colleagues are correct, if social structure is the most powerful variate in explaining differences in brain size when controlling for phylogenetics and body size, then in some ways it is surprising to me. After all, it does not seem that ants have particularly large brains, despite being extremely social and highly successful. Clearly the hymenoptera and other social insects operate on different principles from mammals. Instead of
developing “hive minds,” it seems as if in mammals greater social structure entails greater cognitive structure.

Citation: Susanne Shultz, & Robin Dunbar (2010). Encephalization is not a universal macroevolutionary phenomenon in mammals but is associated with sociality PNAS : 10.1073/pnas.1005246107

(Republished from Discover/GNXP by permission of author or representative)
 
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Kalashpeople_20100312A few days ago I was listening to an interview with a reporter who was kidnapped in the tribal areas of Pakistan (he eventually escaped). Because he was a Westerner he mentioned offhand that to “pass” as a native for his own safety he had his guides claim he was Nuristani when inquiries were made. The Nuristanis are an isolated group in Afghanistan notable for having relatively fair features. His giveaway to his eventual captors was that his accent was clearly not Nuristani, and master logicians that the Taliban are, the inference was made that he was likely a European pretending to be Nuristani.

I thought about this incident when looking over the supplements yesterday of Reconstructing Indian population history. On page 19 note S2 figure 1 includes the Kalash of Pakistan. These are the unconverted cousins of the Nuristanis who were not forcibly brought into the religion of peace in the late 1800s because their region of the Hindu Kush was under British rule, who naturally imposed their late 19th century European value that populations should not be converted by force to a particular religion (Nuristan means “land of light,” whereas before Afghans called it Kafiristan, “land of the unbelievers”). Despite the fair features of the Kalash, which has given rise to rumors that they are the descendants of Alexander the Great’s soldier s, they cluster with Central and South Asian populations, not Europeans. Like the Ainu of Japan it seems superficial similarities to Europeans, at least in relation to the majority population around them, has resulted in an inordinate expectation of total genome exoticism, when in reality a few particular loci are producing the distinctiveness.

Figure 1 from the 2007 paper, Genetic Evidence for the Convergent Evolution of Light Skin in Europeans and East Asians, brings home the point:


skinfig1

The first panel shows a representation of the genetic distance across the genome. Or at least enough to give you a good sense of the phylogenetic relationships. South Asians and Europeans form a clade, as do Native Americans and East Asians. The subsequent panels show Fst values, between population variance, on five genes known at that time to be implicated in between population skin color differences. Note now much different the trees are from the one generated by a large number of loci. Since then more loci have come out of the woodwork, and the peculiar genetic architecture of pigmentation has been rather well characterized. Though most genetic variance may be found within continental races, pigmentation is quite often the exception to this rule. Most of the variance on these loci can be between the races. On SLC24A5 West Africans and Europeans have nearly 100% between population variance. The allele frequencies are disjoint. This shouldn’t be that surprising, skin color is highly heritable, and we already know that there’s a lot of between population difference. So from that one would infer that there would be a lot of genetic variation.

Our skin is our largest organ, and is extremely important as a visual marker of health, age, and identity. The fact that there is so much salient interpopulation difference matters a great deal in the “folk taxonomy” of our species. When considering the relevance of skin color in our taxonomies I thought back to Jared Diamond’s 1994 piece for Discover, Race Without Color:

Regarding hierarchy, traditional classifications that emphasize skin color face unresolvable ambiguities. Anthropology textbooks often recognize five major races: “whites,” “African blacks,” “Mongoloids,” “aboriginal Australians,” and “Khoisans,” each in turn divided into various numbers of sub-races. But there is no agreement on the number and delineation of the sub-races, or even of the major races. Are all five of the major races equally distinctive? Are Nigerians really less different from Xhosas than aboriginal Australians are from both? Should we recognize 3 or 15 sub-races of Mongoloids? These questions have remained unresolved because skin color and other traditional racial criteria are difficult to formulate mathematically.

16 years on I think we can reasonably answer many of Diamond’s questions with phylogenetic trees such as the one to the left. There are five races in the tree by coincidence, though the Khoisans are with Africans, and the Americas has its own branch. Yes, Nigerians are probably less different from the Xhosas than Aboriginal Australians. And I guess this tree implies closer to 15 “subraces” for “Mongoloids.” And with the rise of skin reflectance measures the trait isn’t that difficult to formulate mathematically actually.

But as for the salient phenotypic characteristics which humans use to classify each other, and which will remain important socially and culturally for the near future, it makes absolutely no sense to minimize the critical importance of skin pigmentation. Humans are a very visual species, and the complexion of our largest organ will always be of particular interest. This sort of phenetic classification is not scientifically rigorous, I don’t want cladists to hunt me down, but, it is not an arbitrary cultural construct. We can’t classify people by HLA profiles because we don’t have conscious access to such information (and the idea that we can “smell” HLA profiles is still unproven). Our innate pattern recognition competencies are such that naturally folk taxonomies will start with complexion, and use other characters to refine our categories. Just because something isn’t scientific doesn’t always mean it’s silly or arbitrary.

Image Credit: Outlook India

Addendum: I just realized that Jared Diamond’s article came out at the same time as The History and Geography of Human Genes. A quick consultation of this seminal work would have cleared up some of his questions.

(Republished from Discover/GNXP by permission of author or representative)
 
• Category: Science • Tags: Anthropology, Genetics, Kalash, Race, Taxonomy 
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Razib Khan
About Razib Khan

"I have degrees in biology and biochemistry, a passion for genetics, history, and philosophy, and shrimp is my favorite food. If you want to know more, see the links at http://www.razib.com"