Introduction
... and welcome to the Online Altruism Lab!
This website explores the theoretical foundations of cooperation and altruism in biology, with a particular focus on spacial selection. A series of online simulations and some thought experiments demonstrate that the structuring of a population is sufficient to enable the evolution of altruistic traits.Altruism – A Darwinian Puzzle
In biology, altruism refers to traits in which individuals act in a way that benefits others, but at a cost to themselves. Such traits challenge the traditional view of natural selection, which emphasizes competition and the survival and reproduction of the individual.However, there are countless examples of this in nature. Classic examples are the warning calls in birds, sterile worker castes in social insects, and food sharing among primates. Recent research is uncovering more and more altruistic traits also in microbes, bacteria, fungi, and other organisms (Cremer et. al., 2019).
In each case, the individual performing the altruistic act reduces its own fitness — in extreme cases even to the point of self-sacrifice — while simultaneously promoting the survival or reproductive success of others. Explaining how such altruistic traits evolve has become a central theme in evolutionary biology.
Proposed solutions
Several mechanisms have been proposed for solving the altruism puzzle and the evolution of cooperation, but there is no consensus on their validity. Nowak (Nowak, 2012) distinguishes between the following five mechanisms.
The most widespread approach in biology is kin selection, which relies on the genetic relationship of the cooperating actors (Hamilton, 1964). The main beneficiary of the cooperation is not the individual, but the common "gene" that the relatives share. An individual can therefore even sacrifice its life if this saves enough relatives to make it worthwhile for the shared gene. Only the "inclusive fitness" must show a positive balance.
A different and often seen opposite concept is group selection. The basic idea goes back to Darwin’s assumption that groups compete with each other (Darwin, 1888). This means that groups of individuals, rather than individuals, are the units on which selection acts. A modern version of this, multilevel selection, assumes that selection acts at different levels, such as both the individual and the group level (Sober and Wilson, 1998; Wilson and Wilson, 2008).
Two further solution mechanisms for the altruism puzzle are direct and indirect reciprocity. In the case of direct reciprocity (Trivers, 1971), the costs and benefits of repeated interactions between the actors are extrapolated and analyzed using game theory. In the case of indirect reciprocity, the alleged altruism is rewarded with reputation, which later benefits the actor. Both approaches attempt to eliminate the negative fitness balance of apparent altruism and show that it is in fact not an altruistic trait after all, but serves one's own advantage.
The fifth mechanism for the evolution of cooperation is spatial selection (Nowak, 2012). This is based on the fact that not every individual can interact with every other individual, but that the population is spatially structured and interactions only take place in a limited environment.
Spacial selection and structured populations
The models presented on this website focus on this fifth mechanism. It is a key insights in evolutionary biology that structured populations can strongly support cooperation and even costly altruism (Fletcher and Doebeli, 2009; Fletcher and Zwick, 2004; Killingback et al., 2006; Nowak and May, 1992; Nowak et al., 2010).In a well-mixed population (everyone interacts randomly with everyone else), altruistic behaviors are usually selected against, because defectors (egoists) exploit cooperators (altruists) and gain higher payoffs. But in a structured population, where interactions are localized (e.g., within groups, on a lattice, in spatial neighborhoods, or in social networks), the dynamics can change dramatically.
Since our human minds struggle to understand complex dynamics, the computers should do the hard work for us. For this purpose, I have created the various simulation models presented on this website, all of which can lead to stable altruism based solely on structured populations.
All simulation models were created by me in Netlogo (Wilensky, 1999) and should run in all common browsers. Please contact me if you encounter any problems or errors.
References and further reading
Ackermann, M., Stecher, B., Freed, N.E., Songhet, P., Hardt, W.-D., Doebeli, M., 2008. Self-destructive cooperation mediated by phenotypic noise. Nature 454, 987–990. https://doi.org/10.1038/nature07067
Cremer, J., Melbinger, A., Wienand, K., Henriquez, T., Jung, H., Frey, E., 2019. Cooperation in Microbial Populations: Theory and Experimental Model Systems. Journal of Molecular Biology 431, 4599–4644. https://doi.org/10.1016/j.jmb.2019.09.023
Darwin, C., 1888. The descent of man: and selection in relation to sex. John Murray, Albemarle Street.
Dugatkin, L.A., 2017. The evolution of altruism. Vestn. VOGiS 21, 487–491. https://doi.org/10.18699/VJ17.267
Fletcher, J.A., Doebeli, M., 2009. A simple and general explanation for the evolution of altruism. Proc. R. Soc. B. 276, 13–19. https://doi.org/10.1098/rspb.2008.0829
Fletcher, J.A., Zwick, M., 2004. Strong altruism can evolve in randomly formed groups. Journal of Theoretical Biology 228, 303–313. https://doi.org/10.1016/j.jtbi.2004.01.004
Hamilton, W.D., 1964. The genetical evolution of social behaviour. I. Journal of Theoretical Biology 7, 1–16. https://doi.org/10.1016/0022-5193(64)90038-4
Hardin, G., The Tragedy of the Commons. Science162,1243-1248(1968). DOI:10.1126/science.162.3859.1243
Killingback, T., Bieri, J., Flatt, T., 2006. Evolution in group-structured populations can resolve the tragedy of the commons. Proc. R. Soc. B. 273, 1477–1481. https://doi.org/10.1098/rspb.2006.3476
Kropotkin, P., 1902. Mutual aid: A factor of evolution. McClure Phillips & Co.
Maynard Smith, J., Price, G. The Logic of Animal Conflict. Nature 246, 15–18 (1973). https://doi.org/10.1038/246015a0
Nowak, M.A., 2012. Evolving cooperation. J Theor Biol 299, 1–8. https://doi.org/10.1016/j.jtbi.2012.01.014
Nowak, M. A. & May, R. M., 1992. Evolutionary games and spatial chaos. Nature 359, 826–829. (doi:10.1038/ 359826a0)
Nowak, M. A., Tarnita, C. E., Antal T., 2010. Evolutionary dynamics in structured populations. Phil. Trans. R. Soc. B. 365, 19–30 (doi:10.1098/rstb.2009.0215)
Okasha, S., 2020. Biological Altruism, in: Zalta, E.N. (Ed.), The Stanford Encyclopedia of Philosophy. Metaphysics Research Lab, Stanford University.
Pepper, J.W., 2000. Relatedness in Trait Group Models of Social Evolution. Journal of Theoretical Biology 206, 355–368. https://doi.org/10.1006/jtbi.2000.2132
Price, G.R., 1970. Selection and Covariance. Nature 227, 520–521. https://doi.org/10.1038/227520a0
Sigmund, K., Hauert, C., 2002. Altruism. Current Biology 12, R270–R272. https://doi.org/10.1016/S0960-9822(02)00797-2
Sober, E., Wilson, D.S., 1998. Unto Others: The Evolution and Psychology of Unselfish Behavior, Emersion: Emergent Village Resources for Communities of Faith Series. Harvard University Press.
Steiner, K.F., 2021. The Good, the Bad and the Stochastic: How Living in Groups Innately Supports Cooperation. bioRxiv 2021.02.21.431661; doi: https://doi.org/10.1101/2021.02.21.431661
Steiner, K.F., 2024. Altruism pays off in group-structured populations through probable reciprocity. bioRxiv 2024.01.20.575560. doi: https://doi.org/10.1101/2024.01.20.575560
Trivers, R.L., 1971. The Evolution of Reciprocal Altruism. The Quarterly Review of Biology 46, 35–57. https://doi.org/10.1086/406755
Wilensky, U., 1999. NetLogo. http://ccl.northwestern.edu/netlogo/. Center for Connected Learning and Computer-Based Modeling, Northwestern University. Evanston, IL.
Wilson, D.S., 1975. A theory of group selection. Proc. Natl. Acad. Sci. U.S.A. 72, 143–146. https://doi.org/10.1073/pnas.72.1.143
Wilson, D.S., Wilson, E.O., 2008. Evolution “for the Good of the Group.” American Scientist 96, 380–389.