Tag Archives: Darwin

Between Darwin and Saint Valentine’s day

Yesterday was Darwin’s birthday (and Lincoln’s). Tomorrow is Valentine’s Day. Here’s a post appropriate for either day.

Imagine sex worked like this:

You’ve been feeling bad lately, getting sick a lot. You’re not at your best. You find someone who seems to be in better shape. One thing leads to another and you wind up acquiring body fluids from the other party – and picking up some new genes from them. The new genes help a lot in fighting off infection. You’re feeling better now.

Reproduction? That’s another matter, nothing directly to do with sex. When you reproduce, your offspring will carry all the genes you happen to have at the moment. (Here’s one drawback, according to The Onion.)

Also, I forgot to mention that you’re neither male or female – the gene exchange could have gone in the other direction if you’d both been in the mood. And your partner in the adventure above might not even have been the same species as you. (Just what counts as a species here isn’t well-defined.)

This is more or less how bacteria work out sex. (Joshua Lederberg got the Nobel Prize for figuring this out.) Eukaryotes (you’re one of them) mostly do it differently, combining sex and reproduction. It’s the story you learned in high school about passing on half your genes to a gamete (sex cell), which joins with another gamete to make a new organism.

Most eukaryotes also have two sexes. The best theory we have about why that got started goes like this: Most of the DNA in a eukaryote cell is in the nucleus. But a small fraction is in the mitochondria, little powerhouses outside the nucleus that started out as bacteria, and got domesticated. Imagine that two gametes join together, and combine two sets of mitochondria. There’s a potential conflict here. Suppose your mitochondria have a mutation that lets them clobber your partner’s mitochondria. This is good (evolutionarily speaking) for the winning mitochondria, but very likely to be bad for the cell as a whole. Better for the cell as a whole is if one gamete, acting on instructions from the nucleus, preemptively clobbers all their own mitochondria, so that all the mitochondria come from just the other gamete. This is the beginning of what will eventually lead to a distinction between sperm and eggs, pollen and ovules, male and female. Which means you got all your mitochondrial DNA from your mom, something that will turn out to be important when we look later in the year at geneticists unraveling human prehistory. This is also an example of how selection at one level (within cells) can conflict with selection at another level (between cells). We’ll see such multilevel selection again and again, for example in the evolution of complex human societies.

Sex has to be highly advantageous, although we’re not sure exactly what the advantage is. The general answer is probably that an asexually reproducing organism almost never produces any offspring who have fewer harmful mutations than she has. But a sexually reproducing organism, passing on a random half of her genes to each of her offspring, can have some offspring with fewer harmful mutations, at the cost of having other offspring with more. There are various reasons (Muller’s ratchet, Kondrashov’s hatchet) why this could be evolutionarily advantageous.

In other words, with sexually reproduction, at least some of mum and dad’s kids can be less messed up than their parents; it’s asexually reproducing organisms that really embody Larkin’s dour verse

Man hands on misery to man,

It deepens like a coastal shelf

Get out as early as you can,

And don’t have any kids yourself.

… insofar as, when eukaryote species give up sex, they don’t seem to last long. Dandelions reproduce asexually: based on what we see in other organisms, they probably won’t be around for long, evolutionarily speaking. There’s one mysterious exception, tiny animals called bdelloid rotifers which have been reproducing asexually for tens of millions of years . For readers who are not bdelloid rotifers: Happy Valentine’s Day tomorrow! We’ll have an appropriate evolutionary post up tomorrow

The modern synthesis and the blank slate

August 1938 – November 1943

We’re now dividing time finely enough to include months as well as years.

For most of the later nineteenth century after the publication of On the Origin of Species. biologists were skeptical of Darwin’s proposed mechanism of evolutionary change – natural selection. It was only in the twentieth century that this began to change. When Mendel’s work on heredity was rediscovered in 1900, it was originally seen by many as antithetical to Darwinism. But with the pioneering theoretical work of Fisher, Haldane, and Wright, and the subsequent empirical work of Mayr, Dobzhansky, Simpson, Huxley, Stebbins, and others, Darwin’s theory of natural selection and Mendel’s theory of heredity were combined in what came to be called “the modern synthesis.” Julian Huxley’s book Evolution: The Modern Synthesis marked the coming of age of the theory.

In an earlier post I noted how Lyell’s and Darwin’s embrace of gradualism in explaining the past (as well as George Eliot’s celebration of Dorothea Brooke’s “unhistoric acts” and “hidden life”) had something to do with the political climate in England in the years after the French Revolution and Napoleon. Evolution: The Modern Synthesis was first published in 1942. It’s no surprise that the modern synthesis too was a product of its time, when Nazi Germany and the Soviet Union offered gruesome antithetical demonstrations of how not to think about evolution, genes, and behavior.

Not coincidentally, at the same time that biologists in England and the United States were advancing the modern synthesis, social scientists – cultural anthropologists, behaviorist psychologists – were coming to embrace a strong blank slate view of human nature. (Carl Degler tells the American side of the story in In Search of Human Nature: The Decline and Revival of Darwinism in American Social Thought.) There grew up something amounting to a peace treaty between evolutionary biology and the social sciences, with the two fields agreeing to respect each others’ spheres of influence. Social scientists would leave biology to the biologists, accepting, for example, that neither a good upbringing nor acquired skills can improve your genes. Biologists in turn would largely steer clear of addressing social behavior. For example, the theory of sexual selection, which Darwin developed, and Fisher elaborated, was mostly dropped from the modern synthesis as it matured. Huxley argued (pretty unconvincingly in retrospect) that the elaborate mating dances and ornaments found in so many species were not a product of sexual selection, but merely helped to get individuals to choose the right species of mate. Westermarck’s pioneering work on the evolutionary psychology of incest avoidance and the incest taboo was largely shelved in favor of the shakier theories of Freud and Lévi-Strauss. Even Darwin’s work on emotional expression, which might have seemed fairly anodyne politically, was largely rejected by anthropologists. And the study of prehistory was affected as well.

It was only beginning in the 1960s and 1970s, with the rise of sociobiology, that evolutionary biologists returned to seriously addressing social behavior. Sociobiology: The New Synthesis (1975), by E. O Wilson, made a nod to Huxley in its subtitle. It also announced the end of an intellectual peace treaty, and the opening of an intellectual war that persists up to the present.

Gradualism

1832-1842

Charles Lyell’s great work, Principles of Geology, came out between 1831 and 1833. Lyell advocated an uncompromising uniformitarianism: the same geological forces at work today, causing small changes over the course of lifetimes, were at work in the past, causing massive changes over the course of geological ages. We’ve seen over the course of this blog that uniformitarianism is not a completely reliable guide either to geology or to human history, which have been punctuated often enough by catastrophes – asteroid strikescontinent-scale floodsvolcanic eruptions, and devastating wars and plagues. But the theory is nonetheless at least part of the story of history, and Lyell’s work was deservedly influential.

In 1837 Charles Darwin, a careful reader of Lyell, published a short article entitled On the Formation of Mould. This would eventually led to his last book, The Formation of Vegetable Mould through the Action of Worms. Darwin’s work on soil formation was Lyellianism in miniature. He demonstrated, through a combination of careful reasoning and experiment, that the surface layer of pasture soil is formed by earthworms. “Although the conclusion may appear at first startling, it will be difficult to deny the probability that every particle of earth forming the bed from which the turf in old pasturelands springs, has passed through the intestines of worms.” Reading Darwin on worms you get the feeling he identifies with his humble subjects, gradually remaking the world through their patient industry.

The doctrine of progress through gradual change was appealing for more than just scientific reasons. In the 1830s, English liberals (of whom Darwin was one) were attempting to reform their society gradually, without the violence of the French Revolution, and without turning over politics to a Great Man in the style of Napoleon. (Darwin was also a gradualist with regard to his own work: he came up with the theory of natural selection in 1838, but England at the time wasn’t ready for anything so radical, and he didn’t publish On The Origin of Species for another twenty years.)

George Eliot (Mary Ann Evans), a friend of Darwin’s, set her greatest novel, Middlemarch, around the time of the Reform Act of 1832, which moved England one big step closer to a genuinely representative government. The novel’s heroine, Dorothea Brooke, might in another age have been a famous saint, another Theresa of Avila. In the England of her time she has another fate. Here is the famous conclusion of the novel, a paean to gradualism and the cumulative force of small deeds:

Her full nature … spent itself in channels which had no great name on the earth. But the effect of her being on those around her was incalculably diffusive: for the growing good of the world is partly dependent on unhistoric acts; and that things are not so ill with you and me as they might have been is half owing to the number who lived faithfully a hidden life, and rest in unvisited tombs.

On the Origin of Our Species

88.1 – 83.4 thousand years ago

We saw yesterday that there is recent evidence that Homo sapiens had already made it as far as Saudi Arabia. Eventually, of course, they will replace Neanderthals and Denisovans, with only limited admixture.

One of the Big Questions we run into on Logarithmic History is: To what extent do big changes result from internal causes versus external shocks? Sometimes external shocks are clearly the answer. With the extinction of the dinosaurs, for example, it was a big rock hurtling out of the sky that did the job, not something in dinosaur evolutionary dynamics. Other times the answer is less clear. With the fall of the Roman empire, for example, how much blame falls on external shocks like new infectious diseases or climate change? (A lot, according to Kyle Harper.) How much blame falls on on the historical dynamics of Rome’s dealings with her barbarian neighbors? (A lot, according to Peter Heather.)

We can ask the same question with regard to the evolution of modern humans. Consider the evolution of our capacity for culture. We’ve already mentioned one theory that looks to an external ecological driver, namely the current Ice Age, which (supposedly) put a premium on cultural learning rather than instinct or individual learning. But consider the dynamics of the human expansions Out Of Africa. A currently circulating preprint suggests that these were driven by processes internal to human evolution. Multiple Out Of Africa expansions, and smaller scale expansions, too, would have happened even without climatic instability. Here’s the abstract:

Hominin evolution is characterized by progressive regional differentiation, as well as migration waves, leading to anatomically modern humans that are assumed to have emerged in Africa and spread over the whole world. Why or whether Africa was the source region of modern humans and what caused their spread remains subject of ongoing debate. We present a spatially explicit, stochastic numerical model that includes ongoing mutations, demic diffusion, assortative mating and migration waves. Diffusion and assortative mating alone result in a structured population with relatively homogeneous regions bound by sharp clines. The addition of migration waves results in a power-law distribution of wave areas: for every large wave, many more small waves are expected to occur. This suggests that one or more out-of-Africa migrations would probably have been accompanied by numerous smaller migration waves across the world. The migration waves are considered “spontaneous”, as the current model excludes environmental or other factors. Large waves preferentially emanate from the central areas of large, compact inhabited areas. During the Pleistocene, Africa was the largest such area most of the time, making Africa the statistically most likely origin of anatomically modern humans, without a need to invoke additional environmental or ecological drivers.

A few comments:

First, the model doesn’t explicitly incorporate culture. It does however depend on assortative mating, and of course in our species assortative mating is often a consequence of cultural differences. There are other, complementary models around that explicitly link culture to the population bottlenecks that seem to characterize the evolution of humans (and sperm whales!). I discuss possible implications for the evolution of ethnocentrism here.

Second, in some ways the model hearkens back to a very old idea in evolutionary theory. In On the Origin of Species (Chapter 6, Difficulties on Theory), Darwin wrestles with the question of why the living world is divided into discrete species, with transitional forms relatively rare.

During the process of modification, by which two varieties are perfected into two distinct species, the two which exist in larger numbers … will have a great advantage over the intermediate variety, which exists in smaller numbers in a narrow and intermediate zone. For forms existing in larger numbers will always have a better chance … of presenting further favourable variations for natural selection to seize on, than will the rarer forms which exist in lesser numbers. Hence, the more common forms, in the race for life, will tend to beat and supplant the less common forms, for these will be more slowly modified and improved.

Darwin is arguing for a positive feedback mechanism, a kind of symmetry breaking, whereby more numerous populations evolve and improve more quickly, and drive less numerous, more slowly evolving forms to extinction, thereby creating gaps between species.

There is a widespread idea, put about by Ernst Mayr, that Darwin didn’t have a theory of speciation per se (as opposed to a theory of adaptation by natural selection). But I think Mayr is just wrong; the passage above does offer a theory. Whether this works as a general theory, it does seem to capture something about human evolution from Homo erectus onward. Successive exceptionally well-adapted populations from densely settled areas do a very thorough job of supplanting less common forms, until eventually, in days to come on Logarithmic History, we wind up with a single hugely successful species, Homo sapiens, separated by an enormous gap from its nearest living relatives.

It’s a small world after all

595-563 thousand years ago

The story of human origins is partly a story of Big Things like The Taming of Fire and the The Dawn of Speech. But it’s also the story of some odd byways and quiddities. A nice introduction to some of these is Chip Walter’s book Thumbs, Toes, and Tears: And Other Traits That Make Us Human. (His more recent Last Ape Standing is good too.) Walters considers funny bits of anatomy like our unique big toes and thumbs, and funny bits of behavior like our habits of laughing, weeping, and kissing. Toes and thumbs fossilize, but behaviors can be hard to date, evolutionarily. Presumably these behaviors appeared sometime before modern humans evolved and spread, so let’s pick today’s date. It’s also hard to figure out the exact evolutionary rationale for some of these behaviors. Humor, for example, is not a simple phenomenon: intellectually appreciating a joke, actually finding it funny and enjoying it, and finally laughing, each involve separate areas of the brain.

Laughter, specifically, is a minor human oddity that sheds an interesting light on some big events in human evolution. Robert Provine, a leading laughter researcher, spells out the argument in “Curious Behavior: Yawning, Laughing, Hiccuping, and Beyond.” Chimpanzees have a kind of laugh, a modified vocalized panting synchronized with inhalation and exhalation. Presumably laughter first resulted when panting-during-play evolved into a play signal. But the short bursts of human laughter go further, having freed themselves from synchrony with the inhalation/exhalation cycle. Laughter, in other words, is just one instance of the more general phenomenon of humans having separate controls for vocalization and for respiration. Interestingly, the most prominent examples of complex vocalization – songbirds and some other birds, whales, bats, and humans – are all found in non-quadrupeds. In quadrupeds, breathing is tightly coupled with locomotion: lungs need to be full to stiffen the thorax when the forelimbs hit the ground. Giving up quadrupedalism seems to have allowed for an “adaptive release” in the evolution of vocal abilities in a number of unrelated lineages. So the study of laughter (and other vocalizations) suggests that two key human adaptations – bipedalism and spoken language – are more closely linked than one might have expected.

Another and overlapping set of human particularities involves facial expressions of the emotions. Darwin got a whole book out of this. He concluded (admittedly based on somewhat anecdotal evidence) that different emotional expressions are largely innate. It’s an interesting illustration of his ability to reason from small facts to large conclusions that he also drew a big conclusion about human evolution from this. In Darwin’s day, there were scientists who believed that different human races had evolved from very different prehuman progenitors: one prehuman species giving rise to Europeans, another to Africans, and so on. But Darwin reasoned that the very close similarity in facial expressions (and he had traveled a lot, and witnessed a lot of expressions in a lot of places) and the very similar emotional makeup of humans around the world was evidence that human populations shared a fairly recent common ancestry. Here as in several other cases, a mixture of close reasoning and sheer luck led Darwin to the correct conclusion about evolution long before there was much solid evidence.

Darwin’s work on emotions was neglected for most of the twentieth century by anthropologists favoring a blank slate view of human behavior, but was eventually largely vindicated by a number of researchers, notably Paul Ekman. There is now good evidence for six basic facially expressed emotions: Fear, Disgust, Joy, Anger, Sadness, and Surprise.

If you’re a movie watcher, this list may seem familiar. These emotions (all except for Surprise) are all depicted as little homunculi living inside the head of an 11 year old girl in the animated feature “Inside Out.” Somebody at Pixar Studios knows their Ekman.

insideout

So the sappy song is right: There is just one moon and one golden sun, and a smile means friendship to everyone.

Stonecraft as soulcraft

2.71-2.56 million years ago

Until recently the earliest known stone tools dated back to the Oldowan, 2.6 million years ago, although recently stone tools going back 700,000 years earlier were reported. Stone tool use may be a lot older than we thought.

We now know that tool making is not uniquely human. (For tool use in dinosaurs, see here.) But Oldowan tools – including choppers (below), pounders, and scrapers — go beyond anything chimpanzees, or other animals, do. Kanzi, a bonobo (pygmy chimpanzee, who was also taught to communicate with an artificial set of symbols) learned to use sharp stone flakes for cutting, but never mastered the art of striking a stone core at the proper angle to produce useful sharp flakes. Apparently australopithecines (or maybe early Homo or Kenyanthropus) had taken a step further by 2.6 million years ago (or earlier).

oldowan

Early evolutionary theory developed in tandem with the Industrial Revolution and included an appreciation for the importance of manual labor. Darwin, in The Descent of Man, argued for the central role of toolmaking in human evolution, and, not surprisingly, the same point was echoed by Friedrich Engels in 1876, in his unfinished essay “The Part Played by Labor in the Transition from Ape to Man.” Engels was pushing back against the attitude in most traditional stratified societies that manual labor is low class, while symbolic labor (and/or wielding weapons) is high class.

For example the fingernails on this Chinese scholar advertised that he didn’t work with his hands.

fingernail

Nowadays, a common complaint about the post-industrial economy is that so much education and employment revolves around pushing symbols around that manual labor is relatively devalued. The recent book Shopcraft as Soulcraft: An Inquiry into the Value of Work is a statement of this lament. Maybe today is a good time to celebrate the part played by labor in the transition from ape to man — by making something, or mending something.

But if food is more your thing, the next post will suggest a recipe.

 

Between Darwin and Saint Valentine’s day

Yesterday was Darwin’s birthday (and Lincoln’s). Tomorrow is Valentine’s Day. Here’s a post appropriate for either day.

Imagine sex worked like this:

You’ve been feeling bad lately, getting sick a lot. You’re not at your best. You find someone who seems to be in better shape. One thing leads to another and you wind up acquiring body fluids from the other party – and picking up some new genes from them. The new genes help a lot in fighting off infection. You’re feeling better now.

Reproduction? That’s another matter, nothing directly to do with sex. When you reproduce, your offspring will carry all the genes you happen to have at the moment.

Also, I forgot to mention that you’re neither male or female – the gene exchange could have gone in the other direction if you’d both been in the mood. And your partner in the adventure above might not even have been the same species as you. (Just what counts as a species here isn’t well-defined.)

This is more or less how bacteria work out sex. (Joshua Lederberg got the Nobel Prize for figuring this out.) Eukaryotes (you’re one of them) mostly do it differently, combining sex and reproduction. It’s the story you learned in high school about passing on half your genes to a gamete (sex cell), which joins with another gamete to make a new organism.

Most eukaryotes also have two sexes. The best theory we have about why that got started goes like this: Most of the DNA in a eukaryote cell is in the nucleus. But a small fraction is in the mitochondria, little powerhouses outside the nucleus that started out as bacteria, and got domesticated. Imagine that two gametes join together, and combine two sets of mitochondria. There’s a potential conflict here. Suppose your mitochondria have a mutation that lets them clobber your partner’s mitochondria. This is good (evolutionarily speaking) for the winning mitochondria, but very likely to be bad for the cell as a whole. Better for the cell as a whole is if one gamete, acting on instructions from the nucleus, preemptively clobbers all their own mitochondria, so that all the mitochondria come from just the other gamete. This is the beginning of what will eventually lead to a distinction between sperm and eggs, pollen and ovules, male and female. Which means you got all your mitochondrial DNA from your mom, something that will turn out to be important when we look later in the year at geneticists unraveling human prehistory. This is also an example of how selection at one level (within cells) can conflict with selection at another level (between cells). We’ll see such multilevel selection again and again, for example in the evolution of complex human societies.

Sex has to be highly advantageous, although we’re not sure exactly what the advantage is. The general answer is probably that an asexually reproducing organism almost never produces any offspring who have fewer harmful mutations than she has. But a sexually reproducing organism, passing on a random half of her genes to each of her offspring, can have some offspring with fewer harmful mutations, at the cost of having other offspring with more. There are various reasons (Muller’s ratchet, Kondrashov’s hatchet) why this could be evolutionarily advantageous.

In other words, with sexually reproduction, at least some of mum and dad’s kids can be less messed up than their parents; it’s asexually reproducing organisms that really embody Larkin’s dour verse

Man hands on misery to man,

It deepens like a coastal shelf

Get out as early as you can,

And don’t have any kids yourself.

… insofar as, when eukaryote species give up sex, they don’t seem to last long. Dandelions reproduce asexually: based on what we see in other organisms, they probably won’t be around for long, evolutionarily speaking. There’s one mysterious exception, tiny animals called bdelloid rotifers which have been reproducing asexually for tens of millions of years . For readers who are not bdelloid rotifers: Happy Valentine’s Day tomorrow! We’ll have an appropriate evolutionary post up tomorrow