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E.O. Wilson has passed on.

E.O. Wilson, a Pioneer of Evolutionary Biology, Dies at 92 - The New York Times - "A Harvard professor for 46 years, he was an expert on insects and explored how natural selection and other forces could influence animal behavior. He then applied his research to humans."
As an expert on insects, Dr. Wilson studied the evolution of behavior, exploring how natural selection and other forces could produce something as extraordinarily complex as an ant colony. He then championed this kind of research as a way of making sense of all behavior — including our own.

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Dr. Wilson also became a pioneer in the study of biological diversity, developing a mathematical approach to questions about why different places have different numbers of species. Later in his career, he became one of the world’s leading voices for the protection of endangered wildlife.

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As his parents’ marriage disintegrated, he found solace in forests and tidal pools. “Animals and plants I could count on,” Dr. Wilson wrote in his 1994 memoir, “Naturalist.” “Human relationships were more difficult.”
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After a fish's fin spine poked out one eye, he became very interested in ants, and he would be interested in them for the rest of his life.

At the time, he was also undergoing a spiritual transformation. Raised as a Baptist, he struggled with prayer. During his baptism, he became keenly aware that he felt no transcendence. “And something small somewhere cracked,” Dr. Wilson wrote. He drifted away from the church.

“I had discovered that what I most loved on the planet, which was life on the planet, made sense only in terms of evolution and the idea of natural selection,” Dr. Wilson later told the historian Ullica Segerstrale, “and that this was a far more interesting, richer and more powerful explanation than the teachings of the New Testament.”
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Sociobiology was his groundbreaking work, I just picked it up from the library a few months ago.
 
He did work on the biogeography of ants, and also work on biodiversity, finding that larger islands tend to have more species than smaller ones, and that islands near mainland tend to have more species than more isolated islands. Other biologists extended this work to lakes and forests and the like, and they found that habitat fragmentation creates habitability islands.

He then got into evolution of behavior, and he studied ant societies. Ants communicate by smell, having glands that release various scents.

There is a problem in explaining ant behavior: worker ants don't reproduce. It's only the queens and males that do so.
He found an answer — for a time, at least — in the work of William Hamilton, a British graduate student. Mr. Hamilton argued that biologists needed to focus less on individual animals and more on their genes.

The females in an ant colony were all the daughters of the queen. By caring for the queen’s offspring, they could pass down more of the genes they shared in common.

Mr. Hamilton described “inclusive fitness,” as this concept came to be known, in a 1963 paper. Dr. Wilson took the paper on a long train ride from Boston to Miami. At the start of the journey, he was skeptical; by the end he was convinced.

“I was a convert, and put myself in Hamilton’s hands,” Dr. Wilson wrote.
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Worker insects are thus much like the body cells of multicellular organisms. They don't themselves reproduce, but they assist others in reproducing. This works because of kin selection, assisting others who share one's genes.
 
Then his big 1975 book "Sociobiology: The New Synthesis."

The book got a lot of praise at first.

Dr. Wilson got in trouble for extending sociobiology to humans. He had invited his readers to consider how human nature might be shaped by evolutionary pressures. He warned them that this would not be easy: It would be hard to tease apart the effects of human culture from those of natural selection. Making matters worse, no one at the time had linked any genetic variant to any particular human behavior. “There is a need for a discipline of anthropological genetics,” he wrote.

Nevertheless, Dr. Wilson argued that our species had a propensity to behave in certain ways and form certain social structures. He called that propensity human nature.

Natural selection could help explain psychology, in other words. Human aggression, for example, may have been adaptive for early humans.

“The lesson for man is that personal happiness has very little to do with all this,” he wrote. “It is possible to be unhappy and very adaptive.”
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A lot of that seems like pure hand-waving. We do things that we could not possibly have been selected for, like wear clothing and use written language. If "primitive" people are any guide, than our warm-climate Paleolithic ancestors were nearly naked. But we nowadays wear much more clothing, even in warm weather. Turning to language, spoken language is pretty much a human universal, while written language has been adopted much faster than what natural selection can plausibly produce.
Dr. Wilson’s critics ignored these caveats. In a letter to The New York Review of Books, some denounced sociobiology as an attempt to reinvigorate tired old theories of biological determinism — theories, they claimed, that “provided an important basis for the enactment of sterilization laws and restrictive immigration laws by the United States between 1910 and 1930 and also for the eugenics policies which led to the establishment of gas chambers in Nazi Germany.”

In her book “Defenders of the Truth” (2000), Dr. Segerstrale wrote that Dr. Wilson’s critics had shown “an astounding disregard” for what he had written, arguing that they had used “Sociobiology” as an opportunity to promote their own agendas. When Dr. Wilson attended a 1978 debate about sociobiology, protesters rushed the stage shouting, “Racist Wilson, you can’t hide, we charge you with genocide!” A woman dumped ice water on him, shouting, “Wilson, you are all wet!”

After drying himself off with paper towels, Dr. Wilson went ahead and gave his speech.

In that speech and elsewhere, Dr. Wilson declared that sociobiology offered no excuse for racism or sexism. He dismissed attacks against him as “self-righteous vigilantism.” And he went on to dig even deeper into the evolution of human behavior.
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Despite the importance of his work, he has nevertheless had critics in his home turt: ant behavior.
Among them was Deborah Gordon, a leading expert on ants at Stanford University.

“Wilson’s view of how an ant colony works had every ant genetically programmed to do a certain thing,” Dr. Gordon said in a 2019 interview. “He wanted everybody to do what they were supposed to do without any mess.”

In her own research, Dr. Gordon found that ants can switch from one job to another. And they do not respond to any particular chemical signal like little robots; instead, they will respond differently under different circumstances. “The process is messy,” Dr. Gordon said.

Dr. Wilson vigorously attacked Dr. Gordon’s work, both in print and in person. When Dr. Gordon was at Harvard in the mid-1980s on a fellowship, she recalled Dr. Wilson standing up in the middle of one of her talks to shout his objections. “He really made a lot of effort to keep me from getting a job,” she said.
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Human sociobiology is nowadays called evolutionary psychology, it seems.
Some researchers have tried to construct elaborate evolutionary accounts for how individual genes helped give rise to human nature. But again and again, many of these explanations have proved to be simplistic to the point of misleading. Scientists are a long way from Dr. Wilson’s dream of an evolution-based account of human nature.
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Like everything we do being genetically programmed and directly adaptive.
 
Closer, though, with the advent of epigenetic studies and breathtakingly rapid advances in endocrinology. Sociobiology was audacious in what it was trying to accomplish and concluded ahead of what it could demonstrate empirically, but then so is anthropology as a whole. I was enraptured with Wilson's entymological studies as a youth, then in college learned to be skeptical if not hostile to his social theories, but these days the latter feelings have moderated a bit. We none of us truly know what is going on. As with all true sciences, it is always easier to determine what isn't going on than what is.

E.O. Wilson will be greatly missed.
 
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