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Tag: Evolution
We stand on the shoulders of cultural giants
In reading The cultural niche: Why social learning is essential for human adaptation in PNAS I couldn’t help but think back to a conversation I had with a few old friends in Evanston in 2003. They were graduate students in mathematics at Northwestern, and at one point one of them expressed some serious frustration at the fact that so many of the science and business students in his introductory calculus courses simply wanted to “learn” a disparate set of techniques, rather than understand calculus. The reality of course is that the vast majority of people who ever encounter calculus aim to learn it for reasons of utility, not so that they can grok the fundamental theorem of calculus. With the proliferation of tools such as Mathematica and powerful portable calculators fewer and fewer people are getting their hands dirty with calculus in an analytic sense, and more often see it as simply a “requirement” which they have to pass.
Calculus, and mathematics generally, is a clean and crisp human invention. In the late 17th century Isaac Newton and Gottfried Leibniz originated calculus as we understand it. Later thinkers extended their work. But for the vast majority of humans who have ever learned calculus it is simply a “black box” set of techniques which work rather magically. They did not contribute anything new to the body of knowledge which they drew upon. Mathematics is part of our cultural patrimony, we implicitly stand upon the shoulders of giants without apology. Such is to be human.
Toward evolutionary monism
Is there any substantive difference between natural, sexual, and artificial selection? Or is it just semantic sugar, useful for humans in our own cognitive bookkeeping? I lean toward the latter proposition. To some extent I would think that this is an irrelevant issue, selection is selection, but I have encountered folks who seem surprised at analogies between “artificial” and “natural” selection quite regularly. Of course Charles Darwin famously elided the distinctions across the two categories in his original works in the 19th century (this was later a subject of controversy, insofar as Darwin’s conflation of the properties of artificial and natural selection may have misled him in terms of the weight of factors shaping evolution in the wild).
These are the questions which bubble to the fore of my mind when I encounter reports such as Elizabeth Pennisi’s in Science, On the Trail of Brain Domestication Genes:
Researchers have proposed that bonobos evolved domesticated behavior to encourage group living. By isolating a group of 40 putative brain domestication genes in the prefrontal cortex and comparing their expression in humans versus chimps and bonobos, researchers found that the activity of that gene group in bonobos was clearly “domesticated” compared with chimps, they reported at the Biology of Genomes meeting.
The full piece is gated, so here’s the relevant section in the details:
Saudi Arabia, where monkey became man?
As late as the 1980s it is reputed that prominent Saudi clerics were making the case for geocentrism. Of course presumably most Saudis are not geocentrists, but their religious establishment is so calcified that medieval science still retains some hold upon their imaginations. That’s why I’m very, very, curious about the possibility which is emerging that a critical period of human evolution occurred in the Arabian peninsula. Maju points me to a paper in Quaternary Science Reviews which reports on the discovery of a site in north-central Saudi Arabia, the heartland of the House of Saud, which suggests human occupation ~75,000 years B.P. Middle Paleolithic occupation on a Marine Isotope Stage 5 lakeshore in the Nefud Desert, Saudi Arabia:
Major hydrological variations associated with glacial and interglacial climates in North Africa and the Levant have been related to Middle Paleolithic occupations and dispersals, but suitable archaeological sites to explore such relationships are rare on the Arabian Peninsula. Here we report the discovery of Middle Paleolithic assemblages in the Nefud Desert of northern Arabia associated with stratified deposits dated to 75,000 years ago. The site is located in close proximity to a substantial relict lake and indicates that Middle Paleolithic hominins penetrated deeply into the Arabian Peninsula to inhabit landscapes vegetated by grasses and some trees. Our discovery supports the hypothesis of range expansion by Middle Paleolithic populations into Arabia during the final humid phase of Marine Isotope Stage 5, when environmental conditions were still favorable.
The material describing the site is basically Greek to me, so I had to “hum” my way through. Let’s jump to the final paragraph:
One root for rice
The Pith: There was one rice domestication event somewhere in central China about 10,000 years ago. Probably!
Going by the numbers rice is the real staff of life. Rice is the staple for ~50% of humans alive today. So the science of rice is of major pragmatic importance of all humans (even if a major disease which impacts rice doesn’t result in mass starvation, it will probably generate a price spike for all other staples due to Asian demand). One major issue which I’ve kept track of over the years has been the origin of rice agriculture: was it a parallel multi-hearth origination or a single-hearth event? We know that there were at least two instantiations of agricultural civilization in the world without any cultural diffusion: in the New World and the Old. More likely there were at least several independent hearths in the old world which utilized local wild crops. Wheat and barely in the west, millet and rice in the east, etc. But there is also a model that rice agriculture had two independent origins, in India and China, which gave rise to the indica and japonica strains from the welter of wild rice lineages. Some genetics has supported the model of two hearths by reporting a deep time depth to the last common ancestor of these strains, on the order of ~100,000 years. The implication from phylogenetics is that there were two adaptations of local lines, which later converged in morph due to parallel selection pressures.
The interest in this issue has application to our understanding of human history. Peter Bellwood in First Farmers argues that what L. L. Cavalli-Sforza termed the “great human diasporas” have their roots ultimately in distinctive domestication events. Bellwood goes on to suggest that the contemporary genetic and linguistic patterns of variation we see around us are the products of rapid population growth of these ancient agricultural nuclei (the genome blogger Digonenes works within this framework). The implication then would be that two domestications of rice would imply two population pulses. A single domestication would imply one pulse. Therefore there is a connection here between historical human population genetics and agricultural history & genetics. This stands to reason in that our own species is so parasitic on domestic crops.
Love and arranged marriage
In the wake of yesterday’s review of a paper on heritable variance in trait preferences realized in romantic partners I couldn’t help but be intrigued by this new study out of PLoS ONE, Evolutionary History of Hunter-Gatherer Marriage Practices. It’s actually a pretty thin piece of work in all honesty from what I can tell. They wanted to query ancestral ranges of marriage patterns by mapping the cultural variation in customs onto a phylogenetic tree. To generate that tree they took mtDNA sequences, which to me seems kind of old school. Using the cultural patterns present in living hunter-gatherer groups they presumed they could infer the ancestral state.
So combining these two sources of data they generated this:
They conclude:
The continuing tangling of the human tree
Last summer I made a thoughtless and silly error in relation to a model of human population history when asked by a reader the question: “which population is most distantly related to Africans?” I contended that all non-African populations are equally distant. This is obviously wrong on the face of it if you look at any genetic distance measures. West Eurasians, even those without recent Sub-Saharan African admixture (e.g., North Europeans) are closer than East Eurasians, who are often closer than Oceanians and Amerindians. One explanation I offered is that these latter groups were subject to greater genetic drift through a series of population bottlenecks. In this framework the number of generations until the last common ancestor with Sub-Saharan Africans for all groups outside of Africa should be about the same, but due to evolutionary factors such as more extreme genetic drift or different selective pressures some non-African groups had diverged more from Africans than others in terms of their genetic state. In other words, the most genetically divergent groups in relation to Africans did not diverge any earlier, but simply diverged more rapidly.
Dienekes Pontikos disagreed with such a simple explanation. He argued that admixture or gene flow between Africans and non-African groups since the last common ancestor could explain the differences. I am now of the opinion that Dienekes may have been right. My own confidence in the “serial bottleneck” hypothesis as the primary explanation for the nature of relationships of the phylogenetic tree of human populations is shaky at best. Why my errors of inference?
There were two major issues at work in my misjudgments of the arc of the past and the topology of the present. In the latter instance I saw plenty of phylogenetic trees which illustrated clearly the variation in genetic distance from Africans for various non-African groups. Why didn’t I internalize those visual representations? It was I think the power of the “Out of Africa” (OoA) with replacement paradigm. Even by the summer of 2010 I had come to reject it in its strong form, due to the evidence of admixture with Neanderthals, and rumors of other events which were born out to be true with the publishing of the Denisovan results. But to a first approximation the clean and simple OoA was still looming so large in my mind that I made the incorrect inference, whereby all non-Africans are viewed simply as a branch of Africans without any particular differentiation in relation to their ancestral population. Secondarily, I also was still impacted by the idea that most of the genetic variation you see in the world around us has its roots tens of thousands of years ago. By this, I mean that the phylogeographic patterns of 25,000 years in the past would map on well to the phylogeographic patterns of the present. This assumption is what drove a lot of phylogeography in the early aughts, because the chain of causation could be reversed, and inferences about the past were made from patterns of the present. My own confidence in this model had already been perturbed when I made my errors, but it still held some sort of sway in my head implicitly I believe. It is one thing to move on from old models explicitly, but another thing to remove the furniture from your cognitive basement and attic.
I have moved further from my preconceptions between then and now. It took a while to sink in, but I’m getting there. A cognitive “paradigm shift” if you will. In particular I am more open to the idea of substantive back migration to Africa, as well as secondary migrations out of Africa. A new paper in Genome Research is out which adds some interesting details to this bigger discussion, and seems to weigh in further against my tentative hypothesis that serial bottlenecks and genetic drift can explain variation in distance to Africans of various non-African groups. Human population dispersal “Out of Africa” estimated from linkage disequilibrium and allele frequencies of SNPs:
The Court Jester and the averaging fallacy
The Pith:Climatic and biological evolutionary pressures on an ecosystem complement at different scales. Neither is “dominant,” as that framing is not even wrong.
Yesterday I alluded to the Court Jester hypothesis of evolutionary change, which is often contrasted with the Red Queen hypothesis. The main embarrassment for me as a person who fancies himself a fan of evolutionary process is that I hadn’t ever heard of the Court Jester Hypothesis before yesterday. Therefore I went back to the paper which outlined many of the basic ideas of the model in 2001, Distinguishing the effects of the Red queen and Court Jester on Miocene mammal evolution in the northern Rocky Mountains. To be fair, the hypothesis itself is a tightening of a range of ideas which were long in the air. I did know, for example, about the Turnover-pulse hypothesis. These are all a set of models which emphasize the abiotic selective pressures on life forms, as opposed to the biotic ones. An abiotic pressure would be something like the Younger Dryas cold snap. A biotic pressure might be an exotic invasive species spreading through the landscape.
In my own mind selection is selection, so I didn’t distinguish them too stridently. In fact, most people seem to have abiotic pressures in mind when they conceive of natural selection, so I generally prefer to emphasize the competition and cooperation between and within species. Additionally, it seems that biotic models are more formally tractable and elegantly constructed (I know much of climate change is cyclical, but I assume that “catastrophe” exhibits a poisson distribution?). I generally lack the “thick” knowledge to really make sense of a lot of detailed natural historical treatments, so I probably avoided them because I didn’t think I’d get much out of them. In hindsight, this seems foolish and shortsighted. Rather like economists focusing on equilibrium states because of their ease of modeling when periodic exogenous shocks are a major variable within our real lives.
Now let’s hit the abstract:
Evolutionary pressures, within and without
Foraminifera, Wikimedia Commons
The Pith: The tree if life is nourished by agon, but pruned by the gods. More literally, both interactions between living organisms and the changes in the environment impact the pulsing of speciation and extinction.
No one can be a true “Renaissance Man” today. One has to pick & choose the set of focuses to which one must turn one’s labor to. Life is finite and subject to trade offs. My interest in evolutionary science as a child was triggered by a fascination with paleontology. In particular the megafauna of the Mesozoic and the Cenozoic, dinosaurs and other assorted reptilian lineages as well as the hosts of extinct and exotic mammals which are no more. Obviously I don’t put much time into those older interests at this point, and I’m as much of a civilian when I read Laelaps as you are. More generally when it comes to evolution I focus on the scale of microevolution rather than macroevolution. Evolutionary genetics and the like, rather than paleontology. This is in part because I lean toward a scale independence in evolutionary process, so that the critical issue for me has been to understand the fundamental lowest level dynamics at work. I’m a reductionist.
I am not quite as confident about the ability to extrapolate so easily from evolutionary genetic phenomena upwards in scale as I was in the years past. But let’s set that aside for a moment, and take a stroll through macroevolution. When I speak of natural selection I often emphasize that much of this occurs through competition within a species. I do so because I believe that the ubiquity of this process is often not properly weighted by the public, where there is a focus on competition between species or the influence of exogenous environmental selective pressures. The intra- and inter- species competition dynamic can be bracketed into the unit of selection debate, as opposed to the exogenous shocks of climate and geology. The former are biotic and the latter are abiotic variables which shape the diversity and topology of the tree of life.
A new paper in Science attempts to quantify the effect of these two classes of variables on the evolutionary arc of a particular marine organism over the Cenozoic, roughly the last 65 million years since the extinction of the dinosaurs. Interplay Between Changing Climate and Species’ Ecology Drives Macroevolutionary Dynamics:
Evolution may explain why baby comes early
The Pith: In this post I review a paper which covers the evolutionary dimension of human childbirth. Specifically, the traits and tendencies peculiar to our species, the genes which may underpin those traits and tendencies, and how that may relate to broader public health considerations.
Human babies are special. Unlike the offspring of organisms such as lizards or snakes human babies are exceedingly helpless, and exhibit an incredible amount of neoteny in relation to adults. This is true to some extent for all mammals, but obviously there’s still a difference between a newborn foal and a newborn human. One presumes that the closest analogs to human babies are those of our closest relatives, the “Great Apes.” And certainly the young of chimpanzees exhibit the same element of “cuteness” which is appealing to human adults. Still there is a difference of degree here. As a childophobic friend observed human infants resemble “larvae.” The ultimate and proximate reason for this relative underdevelopment of human newborns is usually attributed to our huge brains, which run up against the limiting factor of the pelvic opening of women. If a human baby developed for much longer through extended gestation then the mortality rates of their mothers during childbirth would rise. Therefore natural selection operated in the direction it could: shortening gestation times. You might say that in some ways then the human newborn is an extra-uterine fetus.
A new paper in PLoS Genetics attempts to fix upon which specific genomic regions might be responsible for this accelerated human gestational clock. An Evolutionary Genomic Approach to Identify Genes Involved in Human Birth Timing:
The flat and the fit
The Pith: In evolution if you want to win in the long run you don’t want all your eggs in one basket, even if that’s the choicest basket. Sh*t happens, and you better have some back up strategies.
Diversity is a major question in evolutionary biology. In particular, why is there so much diversity, so that the tree of life manifests a multitude of morphs? Might there not be some supreme replicator which emerges from the maelstrom to conquer all before it? This is actually the scenario which unfolds in much of science fiction, with monomorphic grey goo eating everything in its path (a more aesthetically differentiated variant of the super-species emerges in Brian W. Aldiss’ Helliconia Winter). As it is, life on earth does not seem to be converging upon an optimum phenotype for all individuals. In contrast, it seems to be going in the opposite direction broadly speaking (thinking on billion year scales), with the shift from the monotony of communal cyanobacteria to the riotous diversity of tropical forest biomes and coral reefs.
There are many ways you might be able to explain this diversity. Temporal and spatial heterogeneity produces perpetually shifting selection pressures, resulting in transient morphs one after the other. Negative frequency dependent selection, whereby the fitness of a phenotype runs up against its own success. This dynamic is one of the drivers of the Red Queen Hypothesis; the evolutionary arms race in some cases witnessing the resurrection of old techniques against which defenses are no longer recalled. Then there is the possibility that the lack of natural selection as an efficacious evolutionary force could allow for the diversification of phenotypes through random drift. Finally, it may simply be that the gusher of mutation is powerful enough that novelty overwhelms selection and drift’s attempt to pare it back.
A new paper in Nature offers up another possibility. It does so by examining the fact that biological diversity remains operative even within a homogenized chemostat. A chemostat in this context refers to a controlled environment where inputs and outputs are balanced to maintain constant equilibrium conditions for a bacterialculture. Therefore, an unbeatable strategy should emerge in this medium perfectly tailored to the environmental constants, resulting in a homogeneous biota to match. Empirically this is not what occurs. So some explanation is warranted.
Metabolic trade-offs and the maintenance of the fittest and the flattest:
Darwin, eh?
At The Intersection Sheril Kirshenbaum posts some rather stark data from Gallup and a Canadian outfit on the differences in attitudes toward evolution between Americans and Canadians. Those Tories are different! The answers seem very similar to those on offer for the General Social Survey’s “CREATION” question. I thought I’d compare Canadians to various American demographics. The question was asked in 2004 of over 1,400 Americans. I find it somewhat ironic in that I think there has been some question as to the Prime Minister of Canada, Stephen Harper, and his attitude toward evolution. Harper is a member of the Evangelical Protestant Christian and Missionary Alliance (and apparently has appointed known Creationists to various government positions, something controversial or notable in Canada). In contrast, Barack Hussein Obama is famously more grounded in evolution than angels.
Space, the final evolutionary frontier
There’s a rather perplexing paper out in PNAS which I stumbled upon today, An evolutionary process that assembles phenotypes through space rather than through time. Perplexing because I wonder if it is almost so obvious as to be boring, in the trivial but true category, or if it points to a rather deep dimensions of evolutionary process which we’ve ignored. The authors themselves offer up that they’re attempting to reintroduce a concept which has long been in the literature, and, they admit that there isn’t much empirical data to test the importance of the dynamic which they’re outlining. For example, after discussing the consequences which might be entailed by the widespread significance of the phenomenon which they’re describe, they note, “We do not have enough data to test this proposition.”
Here’s their abstract:
In classical evolutionary theory, traits evolve because they facilitate organismal survival and/or reproduction. We discuss a different type of evolutionary mechanism that relies upon differential dispersal. Traits that enhance rates of dispersal inevitably accumulate at expanding range edges, and assortative mating between fast-dispersing individuals at the invasion front results in an evolutionary increase in dispersal rates in successive generations. This cumulative process (which we dub “spatial sorting”) generates novel phenotypes that are adept at rapid dispersal, irrespective of how the underlying genes affect an organism’s survival or its reproductive success. Although the concept is not original with us, its revolutionary implications for evolutionary theory have been overlooked. A range of biological phenomena (e.g., acceleration of invasion fronts, insular flightlessness, preadaptation) may have evolved via spatial sorting as well as (or rather than) by natural selection, and this evolutionary mechanism warrants further study.
Where in the world did anatomically modern humans come from?
The Pith: I review a recent paper which argues for a southern African origin of modern humanity. I argue that the statistical inference shouldn’t be trusted as the final word. This paper reinforces previously known facts, but does not add much that both novel and robust.
I have now read the paper which I expressed a touch of skepticism toward yesterday. Do note, I did not dispute the validity of their results. They seem eminently plausible. I was simply skeptical that we could, with any level of robustness, claim that anatomically modern humans arose in southern vs. eastern, or western, Africa. If I had to bet, my rank order would be southern ~ eastern > western. But my confidence in my assessment is very low.
First things first. You should read the whole paper, since someone paid for it to be open access. Second, much props to whoever decided to put their original SNP data online. I’ve already pulled it down, and sent off emails to Zack, David, and Dienekes. There are some northern African populations which allow us to expand beyond the Mozabites, though unfortunately there are only 55,000 SNPs in that case (I haven’t merged the data, so I don’t know how much will remain after combining with HapMap or HGDP data set).
Sweeping through a fly's genome
Credit: Karl Magnacca
The Pith: In this post I review some findings of patterns of natural selection within the Drosophila fruit fly genome. I relate them to very similar findings, though in the opposite direction, in human genomics. Different forms of natural selection and their impact on the structure of the genome are also spotlighted on the course of the review. In particular how specific methods to detect adaptation on the genomic level may be biased by assumptions of classical evolutionary genetic models are explored. Finally, I try and place these details in the broader framework of how best to understand evolutionary process in the “big picture.”
A few days ago I titled a post “The evolution of man is no cartoon”. The reason I titled it such is that as the methods become more refined and our data sets more robust it seems that previously held models of how humans evolved, and evolution’s impact on our genomes, are being refined. Evolutionary genetics at its most elegantly spare can be reduced down to several general parameters. Drift, selection, migration, etc. Exogenous phenomena such as the flux in census size, or environmental variation, has a straightforward relationship to these parameters. But, to some extent the broadest truths are nearly trivial. Down to the brass tacks what are these general assertions telling us? We don’t know yet. We’re in a time of transitions, though not troubles.
Going back to cartoons, starting around 1970 there were a series of debates which hinged around the role of deterministic adaptive forces and random neutral ones in the domain of evolutionary process. You have probably heard terms like “adaptationist,” “ultra-Darwinian,” and “evolution by jerks” thrown around. All great fun, and certainly ripe “hooks” to draw the public in, but ultimately that phase in the scientific discourse seems to have been besides the point. A transient between the age of Theory when there was too little of the empirics, and now the age of Data, when there is too little theory. Biology is a very contingent discipline, and it may be that questions of the power of selection or the relevance of neutral forces will loom large or small dependent upon the particular tip of the tree of life to which the question is being addressed. Evolution may not be a unitary oracle, but rather a cacophony from which we have to construct a harmonious symphony for our own mental sanity. Nature is one, an the joints which we carve out of nature’s wholeness are for our own benefit.
The age of molecular evolution, ushered in by the work on allozymes in the 1960s, was just a preface to the age of genomics. If Stephen Jay Gould and Richard Dawkins were in their prime today I wonder if the complexities of the issues on hand would be too much even for their verbal fluency in terms of formulating a concise quip with which to skewer one’s intellectual antagonists. Complexity does not make fodder for honest quips and barbs. You’re just as liable to inflict a wound upon your own side through clumsiness of rhetoric in the thicket of the data, which fires in all directions.
In any case, on this weblog I may focus on human genomics, but obviously there are other organisms in the cosmos. Because of the nature of scientific funding for reasons of biomedical application humans have now come to the fore, but there is still utility in surveying the full taxonomic landscape. As it happens a paper in PLos Genetics, which I noticed last week, is a perfect complement to the recent work on human selective sweeps. Pervasive Adaptive Protein Evolution Apparent in Diversity Patterns around Amino Acid Substitutions in Drosophila simulans:
After the evolutionary revolution
My post The paradigm is dead, long live the paradigm! expressed to some extent my befuddlement at the current state of human evolutionary genetics and paleoanthropology. After the review of the paper of possible elevated admixture with Neandertals on the dystrophin locus a friend emailed, “Remember when we thought everything would be so simple once we could finally see this stuff?” Indeed I do remember. The fact that things aren’t simple is very exhilarating, but it is also a major quash on theoretical clarity. Science is after all not a collection of facts, but it is in part facts which one can sieve through a analytic framework.
In hindsight with the relative robustness of ancient DNA results we can make some assessments about the role of human bias within particular heuristic frameworks over the past generation. From the mid-1980s up until 2000 it was victory after victory for the Out-of-Africa with total replacement model. The rise of mtDNA and Y chromosomal lineage studies seemed to buttress the idea of common descent from neo-Africans within the last 100-200,000 years for all human populations. There wasn’t much of a perturbation from this march toward paradigm ascendancy in the aughts, except that there were now also now a trickle of papers which claimed to phylogenetic “long branches” in the human genome. The 2006 Evans et al. paper, Evidence that the adaptive allele of the brain size gene microcephalin introgressed into Homo sapiens from an archaic Homo lineage, was probably the one that made the biggest media splash. But these were inferences. Subsequent analysis of the draft Neandertal genome seems to suggest that in fact the microcephalin allele in question did not introgress.
Case closed? Obviously not. Now we’re in a different era. The Evans et al. paper may have wrong in the specifics, but its general framework seems to likely have been validated: there are genetic lineages in the modern human genome which are not derived from the neo-Africans. But, let us remember that the overwhelming majority of the human genome is neo-African. A reasonable interval for non-Africans is 90-99% neo-African. But, a non-trivial minority has introgressed or admixed from other lineages. Out-of-Africa is mostly correct, but in some ways so is Multiregionalism. But how do we describe this? “Weighted multiregionalism”? “Mostly Out-of-Africa?” The old terms were nice because they were punchy and precise. If you look at Multiregionalism or Out-of-Africa in Wikipedia the newest results are noted, but it doesn’t seem that they’ve been integrated into the analytic narrative. Yet.
More bad mutations = greater fitness
Does the chart above strike you as strange? What it shows is that the mean fitness of a population drops as you increase the rate of deleterious mutation (many more mutations are deleterious than favorable)…but at some point the fitness of the population bounces back, despite (or perhaps because of?) the deleterious mutations! This would seem, to me, an illustration of bizzaro-world evolution. Worse is better! More is less! Deleterious is favorable? By definition deleterious isn’t favorable, so one would have to back up and check one’s premises.
And yet this seems just what a new paper in PLoS ONE is reporting. Purging Deleterious Mutations under Self Fertilization: Paradoxical Recovery in Fitness with Increasing Mutation Rate in Caenorhabditis elegans:
Compensatory mutations can be more frequent under high mutation rates and may alleviate a portion of the fitness lost due to the accumulation of deleterious mutations through epistatic interactions with deleterious mutations. The prolonged maintenance of tightly linked compensatory and deleterious mutations facilitated by self-fertilization may be responsible for the fitness increase as linkage disequilibrium between the compensatory and deleterious mutations preserves their epistatic interaction.
Got that? OK, you probably need some background first….
Men at work: hoes, ploughs, and steel
Ancient Egyptian farmer ploughing a field
Recently several weblogs have pointed to a new working paper on the role of plough-based agriculture vs. hoe-based agriculture in shaping cultural expectations about male and female labor force participation specifically, and the differentiation of gender roles more generally. My first reaction was: “doesn’t everyone know this already?” I am a cursory reader of the anthropological literature and the assumption that a shift from relatively extensive hoe-based agriculture (e.g., slash & burn, gardening, etc.) to a more intensive plough-based mode of production seems to suffuse the literature. Before touching on the major points in the paper itself I did a quick literature search, on the order of five minutes, and found something from 1928 which already assumed the major parameters which are now being mooted today, The Division of Work According to Sex in African Hoe Culture. I read the whole paper, and it remains surprisingly relevant (though some of the terminology and frameworks are a bit dated naturally). Here’s a selection:
Eduard Han, to whom the ethnological study of economics owes a considerable number of important discoveries which have been published repeatedly and in varying forms, seems to have paid scarcely enough attention to the good work of the scholars who preceded him in the fight for the recognition of the outstanding position of women in the lower forms of soil cultivation. Steinmetz and quite recently Koppers,’ have pointed out that Buckland already attributed to the female sex the invention of the most ancient method of soil cultivation, or hoe culture…Here we find, in particular,a clearer statement of the arguments of Grosse, Bachofen, and others about the connexion of matriarchal society and lower forms of soil cultivation. Matriarchy and hoe culture are assigned to definite chronologically determined stages of civilization (older forms of the so-called ‘two class culture’, and later ones of ‘bow culture’). Koppers, of the Austrian branch associates matriarchy and hoe culture with these two civilizations….
It is not our business here to study in detail the researcheson the zones of culture,which may be regardedas successfulup to a certain point, though we shall have to refer to them incidentally. It suffices to state that a connexion between woman and hoe culture, nay more, between that social system where the woman rules, matriarchal society, and primitive soil cultivation is universally acknowledged to exist.
Ignore some of the terms and concepts which might seem loaded or outmoded today. Rather, observe that in 1928 a distinction between hoe-based and plough-based agriculture was widely accepted in terms of the cultural consequences. Why? Just take a look at an old-fashioned plough vs. hoe (at least old-fashioned compared to the sort of mechanized devices you can find in catalogs today):
The inevitable social brain
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.
Sons of the conquerors: the story of India?
The past ten years has obviously been very active in the area of human genomics, but in the domain of South Asian genetic relationships in a world wide context it has seen veritable revolutions and counter-revolutions. The final outlines are still to be determined. In the mid-1990s the conventional wisdom was that South Asians were a branch of a broader West Eurasian cluster of peoples, albeit more distant from the core Middle Eastern-North-African-European-Caucasian clade. The older physical anthropological literature would have asserted that South Asians were predominantly Caucasoid, but with a Australoid element admixed in at varying proportions as a function of geography and caste. To put it more concretely, and I think accurately, a large degree of South Asian physical variety can be defined along the spectrum between A. R. Rahman and Nawaz Sharif. The regional and caste truisms are only correlations. Subrahmanyan Chandrasekhar was a Tamil Brahmin, but experienced anti-black racism in the United States. I think that is reasonable in light of his appearance.
This rough & ready mainstream understanding, supporting by classical genetic markers, was overturned in the early years of the 21st century. One line of thought argued that South Asians were much more distinctive from the broader Western Eurasian cluster of peoples. Representative of this body of work is a paper like The genetic heritage of the earliest settlers persists both in Indian tribal and caste populations. These researchers tended to start with the female lineages, mtDNA, and then supplement that with Y lineages, the paternal descent. A separate line of evidence, generally drawn from Y chromosomal results, indicated that there were deep connections between the people of India and those of Central Eurasia, in particular via the R1a haplogroup. Additionally, one aspect of the first set of results which was very surprising was that it actually placed South Asians closer to East, not West, Eurasians. But by the end of the aughts the uniparental studies had been supplemented by a range of results produced from SNP-chips, which looked at hundreds of thousands of genetic variants. These studies seemed to support the older view of South Asians being closer to West Eurasians than East Eurasians. Finally last year a paper came out which posited that almost all South Asian populations were actually an ancient stabilized hybrid between two groups, a European-like population, “Ancient North Indians” (ANI), and another group which is no longer present in unadmixed form, “Ancient South Indians” (ASI), of whom the Andaman Islanders are distant relatives. Though there was a slight bias toward ANI as a whole, the fraction of ASI increased as one went southeast, and down the caste ladder. The distinctive “South Asian” ancestral group in other words then may actually be conceived of as a compound of these two elements; an admixture of the native substrate against a European-like genetic background.
'dem bones tell strange tales
There is a new paper in PNAS on remains from China which re-order and muddle our understanding of the emergence of anatomical and behavioral modernity in Eurasia. Human remains from Zhirendong, South China, and modern human emergence in East Asia:
The 2007 discovery of fragmentary human remains (two molars and an anterior mandible) at Zhirendong (Zhiren Cave) in South China provides insight in the processes involved in the establishment of modern humans in eastern Eurasia. The human remains are securely dated by U-series on overlying flowstones and a rich associated faunal sample to the initial Late Pleistocene, >100 kya. As such, they are the oldest modern human fossils in East Asia and predate by >60,000 y the oldest previously known modern human remains in the region. The Zhiren 3 mandible in particular presents derived modern human anterior symphyseal morphology, with a projecting tuber symphyseos, distinct mental fossae, modest lateral tubercles, and a vertical symphysis; it is separate from any known late archaic human mandible. However, it also exhibits a lingual symphyseal morphology and corpus robustness that place it close to later Pleistocene archaic humans. The age and morphology of the Zhiren Cave human remains support a modern human emergence scenario for East Asia involving dispersal with assimilation or populational continuity with gene flow. It also places the Late Pleistocene Asian emergence of modern humans in a pre-Upper Paleolithic context and raises issues concerning the long-term Late Pleistocene coexistence of late archaic and early modern humans across Eurasia.
I read the paper, and I really didn’t understand anything between the introduction and discussion. Mostly because it was a detailed exploration of anatomical details, and I’ve never taken an anatomy class. I basically rely on people like John Hawks to tell me what’s going on in that domain. He hasn’t blogged the paper (well, as of this writing), but he did give an assessment to National Geographic:
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