Cooperation: How to Model It, How to Foster It, And How It Might Have Emerged

In a world of self-interested individuals, does cooperation makes sense? The course will explore the phenomenon of cooperation: how it might arise under a set of precise, concrete assumptions about the individuals of a society, and the mechanisms involved in its emergence and continued viability.


Quick reminder that the November 23 lecture will be on Zoom, at this link. Join at 14:15, or a minute or two earlier!


  1. Logistics [pdf, Canva]

  2. Cooperation In The Field [pdf, Canva]

  3. Game Theory 101: Stags, Prisoners and Equilibria [pdf, Canva]

  4. Iterated Games [pdf, Canva]

Research Questions For the Final Assignment

  1. Create a schematic model of the game theory of cancer. Discuss the assumptions behind it, and the consequences for the host organism. See [4, 6].
  2. Discuss the game theory of specialization based on caste in eusocial insects (e.g., bees or ants). See [2].
  3. How does Tit-For-Tat work in a repeated Prisoner’s Dilemma? Is it an equilibrium?
  4. Reciprocity in the natural world. See, for instance, [1, 3].
  5. How can we study altruism in people using economic games? And how do people perform in these games?


  1. Trivers, R. L. (1971). The Evolution of Reciprocal Altruism. The Quarterly Review of Biology, 46(1), 35–57.
  2. Wilkinson, G. S. (1984). Reciprocal food sharing in the vampire bat. Nature, 308(5955), 181–184.
  3. Milinski, M. (1987). TIT FOR TAT in sticklebacks and the evolution of cooperation. Nature, 325(6103), 433–435.
  4. Ridley, M. (1997). The Origins of Virtue: Human Instincts and the Evolution of Cooperation. Penguin.
  5. Roberts, G., & Sherratt, T. N. (1998). Development of cooperative relationships through increasing investment. Nature, 394(6689), 175–179.
  6. Sachs, J. L., Mueller, U. G., Wilcox, T. P., & Bull, J. J. (2004). The evolution of cooperation. The Quarterly Review of Biology, 79(2), 135–160.
  7. Nowak, M. A., & Sigmund, K. (2005). Evolution of indirect reciprocity. Nature, 437(7063), 1291–1298.
  8. Ratnieks, F. L. W., Foster, K. R., & Wenseleers, T. (2006). Conflict resolution in insect societies. Annual Review of Entomology, 51, 581–608.
  9. Axelrod, R. (2006). The Evolution of Cooperation: Revised Edition. Basic Books.
  10. Nowak, M. A. (2006). Five rules for the evolution of cooperation. Science, 314(5805), 1560–1563.
  11. Gächter, S., Herrmann, B., & Thöni, C. (2010). Culture and cooperation. Philosophical Transactions of the Royal Society of London, 365(1553), 2651–2661.
  12. Nowak, M., & Highfield, R. (2011). SuperCooperators: Altruism, Evolution, and Why We Need Each Other to Succeed. Simon and Schuster.
  13. Rand, D. G., & Nowak, M. A. (2013). Human cooperation. Trends in Cognitive Sciences, 17(8), 413–425.
  14. Hummert, S., Bohl, K., Basanta, D., Deutsch, A., Werner, S., Theissen, G., Schroeter, A., & Schuster, S. (2014). Evolutionary game theory: cells as players. Molecular bioSystems, 10(12), 3044–3065.
  15. Aktipis, C. A., Boddy, A. M., Jansen, G., Hibner, U., Hochberg, M. E., Maley, C. C., & Wilkinson, G. S. (2015). Cancer across the tree of life: cooperation and cheating in multicellularity. Philosophical Transactions of the Royal Society of London, 370(1673).
  16. Carter, G. G., Farine, D. R., Crisp, R. J., Vrtilek, J. K., Ripperger, S. P., & Page, R. A. (2020). Development of New Food-Sharing Relationships in Vampire Bats. Current Biology, 30(7), 1275–1279.
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