Whether competition or cooperation is a basic factor in biological interactions between biological individuals or any economic or social interactions between human individuals has been a continuing controversy since Darwin. This is the first of a series of posts on cooperation. The first post will be about the human side of this controversy and all the colorful and tragic characters involved. The second will be about the theoretical side,
especially group selection and mutualism. The third will look at cooperative breeding in vertebrates (mostly birds) and the empirical evidence that has been gathered.
On November 24, 1859, Darwin’s On the Origin of the Species was published. It is hard to imagine how this one book changed intellectual thought. For one thing, this was the point where biology became a science, where the concept of God as a static creator was replaced with the idea of creation and change through biological entities interacting with their environment.
Marx and Engels were both aware of and supportive of Darwin’s theories. Marx attended one of Darwin’s lectures at Oxford and had him sign a copy of Species (24). Marx and Engels emphasized how the theory removed religion from the science of biology by showing that evolution is a system with no direction or purpose, as opposed to teleology, evolution having a direction or purpose, now called “Design” by the creationists. So they merged Darwinism into their anti-religious views and framed evolution into the materialist dialectic and the struggle between workers and bosses. (3) Others used Darwin for various ideologies including apologies for unrestrained capitalism which came to be called Social Darwinism, the “science” of eugenics (sterilizing the unfit), and various racial superiority theories. It was the philosopher and founder of sociology, Herbert Spencer who first coined the term “survival of the fittest” and although Spencer has been brought back in favor as a much more nuanced thinker, his ideas were used by industrialists like Andrew Carnegie as an excuse for unbridled capitalism. (4) (5) (6) Darwin’s cousin, Sir Francis Galton, was the founder of modern statistics and a major proponent of eugenics and the use of selective breeding to improve the human race. (7) One of his students, Roland Fischer, went on to play an important role in the great synthesis of Darwinian evolution and genetics that took place during the 1920s and 1930s. In the United States, poor women, mostly black or native American were sterilized up until the early 1970s, and in India in 1976, during a 21-month state of emergency, 6.2 million men were forcibly sterilized and women are still being sterilized to this day. The idea that the struggle for existence resulted in fierce competition between individuals of different species and between individuals of the same species became dominant in biological thought, especially in the English-speaking world.
Two aspects of animal life impressed me most during the journeys which I made in my youth in Eastern Siberia and Northern Manchuria. One of them was the extreme severity of the struggle for existence which most species of animals have to carry on against an inclement Nature; the enormous destruction of life which periodically results from natural agencies; and the consequent paucity of life over the vast territory which fell under my observation. And the other was, that even in those few spots where animal life teemed in abundance, I failed to find — although I was eagerly looking for it — that bitter struggle for the means of existence, among animals belonging to the same species, which was considered by most Darwinists (though not always by Darwin himself) as the dominant characteristic of the struggle for life, and the main factor of evolution. (2)
Thus starts the introduction of Pëtr Kropotkin’s essay, Mutual Aid A Factor of Evolution, written in 1902. Besides being a major thinker and proponent of anarchism during the Russian Civil War, Kropotkin was also a naturalist who came to a very different conclusion than Darwin while traveling through northeastern Asia. Kropotkin called this Mutual Aid and extended this idea to humans and his theories on Anarchy. In biology, this drives the concept of mutualism, the study of cooperative interactions between different species. A major researcher in this field, Dr. Judith Bronstein, is at the University of Arizona and edited a book, Mutualism, in 2018. (8) Darwin did not fail to notice the factor of cooperation in the animal world and especially among humans. He struggled with this question. Why do individuals cooperate and how does cooperation evolve under natural selection?
Now, if some one man in a tribe, more sagacious than the others, invented a new snare or weapon, or other means of attack or defense, the plainest self-interest, without the assistance of much reasoning power, would prompt the other members to imitate him; and all would thus profit. (1)
Therefore, it hardly seems probable that the number of men gifted with such virtues, or that the standard of their excellence, could be increased through natural selection, that is, by the survival of the fittest; for we are not here speaking of one tribe being victorious over another. (1)
Here Darwin seems to be talking about is what is now called group selection, a hierarchy of selection at the group rather than the individual level. The synthesis of Darwinism and genetics created an evolutionary theory based on genetics with the competition of individual genes mediating adaptation and evolutionary change. Despite the great success of the Synthesis, there were still questions from developmental biologists, from those that studied the fossil record, insect biologists, and those that studied social behavior. In 1962 the first book espousing group selection was published by VC Wynne-Edwards based on his studies of bird behavior during the 1950s.
The first theoretical breakthroughs to the understanding of cooperation from Darwinian genetics came during the 1960s and 1970s with kin selection, derived through the work of WD Hamilton (10)(11), George R Price (12)(13), and John Maynard Smith (13). Hamilton’s rule is a simple inequality based on the idea that altruism (cooperation) between individuals comes from how closely related they are since by helping relatives they are also helping their own genes. This rule was derived from the Price equation, a much richer statistical theory of cooperation based on the work of Roland Fisher and the evolutionary game theory created by Price and Smith. One of the conclusions of all this was that cooperation was impossible between unrelated individuals. George Price became despondent over the implications of his discovery. He started giving away all his money and eventually became homeless. On January 6th, 1975, he slit his throat with nail scissors. He was buried in a paupers grave attended only by his old friends, John Maynard Smith and WD Hamilton.
In 1975 the great Harvard biologist and expert on eusociality in insects, EO Wilson published Sociobiology: The New Synthesis (15) which enshrined Hamilton’s Rule as an integral part of the study of animal behavior. Sociobiology struck a raw cord in some because the idea that genes controlled behavior looked like a revival of eugenics and indeed there was a revival of social Darwinism in the 1980s.
"They are casting their problems at society. And, you know, there's no such thing as society. There are individual men and women and there are families. And no government can do anything except through people, and people must look after themselves first. It is our duty to look after ourselves and then, also, to look after our neighbours." Margaret Thatcher 1987 (14)
Whether Thatcher (or her speechwriters) knew about Hamilton’s rule is unknown but the social Darwinism perspective stands out.
Game theory started in economics to model interactions between two individuals based upon a matrix of payoffs. After major theoretical breakthroughs by John Forbes Nash (16), it was developed in a biological sense by Smith and Price (13). Economics is not called a dismal science for no reason and game theory started out pretty dismal. Models like The Prisoner’s Dilemma and The Tragedy of the Commons showed that cooperation was impossible. In 1981, The Evolution of Cooperation was published by Robert Axelrod and WD Hamilton (17). In it, they showed that the Prisoner’s Dilemma had a solution if it was played again and again over time. Thus a dynamic individualistic game had a cooperative solution. The field of evolutionary dynamics was born.
EO Wilson had become increasingly dissatisfied with the progress in sociobiology and increasingly drawn to group selection. He began working with a prominent group selectionist: David Sloan Wilson (no relation.) In 2007 they co-published: Rethinking the Theoretical Foundation of Sociobiology. (18) This was at a time when group theory was considered both equivalent to and harder to formulate than kin theory. In 2010 he had found his theoretical backing and published The Evolution of Eusociality (19) with Martin Nowak and his postdoc from Yale, Corina Tarnita. In it, they stated that Hamilton’s Rule was just one of five different paths to cooperation. This created a firestorm in the biology world that lasts to this day.
Dr. Martin Nowak is a mathematician who studies game theory and evolutionary dynamics and is now at Harvard University. Born in Austria, he studied at the University of Vienna and later went to Oxford to work with Robert May. He started the first research program in Theoretical Biology at the Institute for Advanced Study at Princeton in 1998. He became the director of the Center for Evolutionary Dynamics at Harvard in 2003. (see FOOTNOTE)
In 2017 Nowak and Wilson published another, even more, a controversial paper stating that Hamilton’s rule is basically of no worth in biology (21). So the debate continues with group selection still a minority opinion among biologists. In future posts, I’ll look more at the theoretical controversy and the empirical evidence with cooperatively breeding birds.
FOOTNOTE:
There have been recent problems with Dr. Nowak’s career. This is exemplified by this image from the end of one of his papers.
(25)
There are two odd sources of money here. One is the John Templeton Foundation (20). John Templeton was a British-American investor who started a foundation to explore the intersection of religion, science, and philosophy and now has a 2.5 billion dollar endowment. They fund scientists and philosophers around the world. They have been accused of a stealth promotion of religion in the sciences. I found no references to religion in any of Nowak’s papers except for one journal about how religion facilitates cooperation in humans (22). I looked at the Templeton web site and found two grants to Nowak in 2016 and 2019 totally $3.5 million. Personally, Nowak’s religious beliefs are his own business as long as it doesn’t affect his duties as a mathematician and scientist. More troubling has been his relationship with the last donor on the list, Jeffery Epstein. Of the 10 million dollars that Epstein gave to Harvard, 6.5 million went to Nowak’s Center for Evolutionary Dynamics. Epstein even had an office there and visited it at least 40 times, often accompanied by young female “assistants.” An investigation by Harvard in 2020 contends that Nowak took money from Epstein even after Epstein was convicted as a sex offender in Florida and was given a suspiciously lenient sentence. In April of 2020 Nowak was placed on administrative leave by Harvard and as far as I know, has not been reinstated (23). To my knowledge, none of Epstein’s victims have ever accused Nowak of anything but his career is forever tainted by this association. In 2019 Nowak published 13 papers, in 2020 he published only 3.
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Kropotkin, Pëtr. Mutual Aid A Factor of Evolution, 1902. https://theanarchistlibrary.org/library/petr-kropotkin-mutual-aid-a-factor-of-evolution.
Angus, Ian. “Marx and Engels...and Darwin?” International Socialist Review. Accessed February 5, 2021. /issue/65/marx-and-engelsand-darwin.
Leonard, Thomas C. “Origins of the Myth of Social Darwinism: The Ambiguous Legacy of Richard Hofstadter’s Social Darwinism in American Thought.” Journal of Economic Behavior & Organization 71, no. 1 (July 2009): 37–51. https://doi.org/10.1016/j.jebo.2007.11.004.
Weinstein, David. “Herbert Spencer.” In The Stanford Encyclopedia of Philosophy, edited by Edward N. Zalta, Fall 2019. Metaphysics Research Lab, Stanford University, 2019. https://plato.stanford.edu/archives/fall2019/entries/spencer/.
Hofstadter, Richard. Social Darwinism in American Thought, 1955. https://www.penguinrandomhouse.com/books/204052/social-darwinism-in-american-thought-by-richard-hofstadter/.
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Bronstein, Judith L. Mutualism. Oxford University Press, 2018. https://global.oup.com/academic/product/mutualism-9780199675654?cc=us&lang=en&.
Wynne-Edwards, Vero Copner. Animal Dispersion in Relation to Social Behaviour. Oliver and Boyd, 1962.
Hamilton, W.D. “The Genetical Evolution of Social Behaviour. I.” Journal of Theoretical Biology 7, no. 1 (July 1964): 1–16. https://doi.org/10.1016/0022-5193(64)90038-4.
Hamilton, W.D. “The Genetical Evolution of Social Behaviour. II.” Journal of Theoretical Biology 7, no. 1 (July 1964): 17–52. https://doi.org/10.1016/0022-5193(64)90039-6.
Price, George R. “Fisher’s ’Fundamental Theorem’ Made Clear.” Annals of Human Genetics 36, no. 2 (November 1972): 129–40. https://doi.org/10.1111/j.1469-1809.1972.tb00764.x.
Smith John Maynard, and George R. Price. “The Logic of Animal Conflict.” Nature 246 (November 2, 1973).
Staff, Guardian. “Margaret Thatcher: A Life in Quotes.” the Guardian, April 8, 2013. http://www.theguardian.com/politics/2013/apr/08/margaret-thatcher-quotes.
Wilson, E. O. Sociobiology - The New Synthesis. Harvard University Press, 1975.
Nash, John F. “Equilibrium Points in N-Person Games.” Proceedings of the National Academy of Sciences of the United States of America 36, no. 1 (1950): 48–49.
Axelrod, R., and W. D. Hamilton. “The Evolution of Cooperation.” Science 211, no. 4489 (March 27, 1981): 1390–96. https://doi.org/10.1126/science.7466396.
Wilson, David Sloan, and Edward O. Wilson. “Rethinking the Theoretical Foundation of Sociobiology.” The Quarterly Review of Biology 82, no. 4 (December 1, 2007): 327–48. https://doi.org/10.1086/522809.
Nowak, Martin A., Corina E. Tarnita, and Edward O. Wilson. “The Evolution of Eusociality.” Nature 466, no. 7310 (August 26, 2010): 1057–62. https://doi.org/10.1038/nature09205.
Templeton, John. “Home - John Templeton Foundation.” Accessed January 29, 2021. https://www.templeton.org/.
Nowak, Martin A., Alex McAvoy, Benjamin Allen, and Edward O. Wilson. “The General Form of Hamilton’s Rule Makes No Predictions and Cannot Be Tested Empirically.” Proceedings of the National Academy of Sciences 114, no. 22 (May 30, 2017): 5665–70. https://doi.org/10.1073/pnas.1701805114.
Rand, David G., Anna Dreber, Omar S. Haque, Rob J. Kane, Martin A. Nowak, and Sarah Coakley. “Religious Motivations for Cooperation: An Experimental Investigation Using Explicit Primes.” Religion, Brain & Behavior 4, no. 1 (January 2, 2014): 31–48. https://doi.org/10.1080/2153599X.2013.775664.
Rosenberg, John S. “Jeffrey Epstein’s Extensive Harvard Reach.” Harvard Magazine, June 8, 2020. https://harvardmagazine.com/2020/07/jhj-epstein-extensive-reach.
Colp, Ralph. “The Contacts Between Karl Marx and Charles Darwin.” Journal of the History of Ideas 35, no. 2 (1974): 329–38. https://doi.org/10.2307/2708767.
Nowak, Martin A. “Five Rules for the Evolution of Cooperation.” Science (New York, N.y.) 314, no. 5805 (December 8, 2006): 1560–63. https://doi.org/10.1126/science.1133755.
“Cooperation: A Divine Virtue for Success- The New Indian Express.” Accessed April 24, 2021. http://cms.newindianexpress.com/cities/chennai/2019/mar/13/cooperation-a-divine-virtue-for-success-1950391.html.
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