Evolution of Organismic Systems

Peter Hammerstein

is a founding professor of the ITB. Though officially retired, he is an active member of the ITB and an external professor of the Santa Fe Institute in New Mexico. A biographical sketch can be found at the bottom of this page.

Research interests

Evolutionary theory across the life sciences: Conflict and cooperation between and within organisms, economics in nature, evolutionary game theory, biological markets, evolution of decision mechanisms, endosymbiont-host interactions, evolutionary medicine/anthropology


Hammerstein, P. & Stevens, J.R., eds. (2012). Evolution and the Mechanisms of Decision Making. Cambridge, MA: MIT Press.

Hammerstein, P., ed. (2003). Genetic and Cultural Evolution of Cooperation. Cambridge, MA: MIT Press.

Noë, R., van Hooff, J.A.R.A.M. & Hammerstein, P., eds. (2001). Economics in Nature. Cambridge: Cambridge University Press.

Albers, W., Güth, W., Hammerstein, P., Moldovanu, B. & van Damme, E., eds. (1997). Understanding Strategic Interaction - Essays in Honor of Reinhard Selten. Berlin: Springer-Verlag.

Selected publications

More publications on Google Scholar.

Zug, R. & Hammerstein, P. (2017) Evolution of reproductive parasites with direct fitness benefits. Heredity, doi:10.1038/s41437-017-0022-5.

Gebiola, M., Kelly, S.E., Velten, V., Zug, R., Hammerstein, P., Giorgini, M., Hunter, M.S. (2017). Reproductive interference and fecundity affect competitive interactions of sibling species with low mating barriers: Experimental and theoretical evidence. Heredity, 119, 438-446.

Hammerstein, P. & Noë, R. (2016). Biological trade and markets. Philosophical Transactions of the Royal Society B, 371, 20150101.

Gebiola, M., Kelly, S., Hammerstein, P., Giorgini, M., Hunter, M.S. (2016). “Darwin’s corollary” and cytoplasmic incompatibility induced by Cardinium may contribute to speciation in Encarsia wasps (Hymenooptera: Aphelinidae). Evolution, doi:10.1111/evo.13037.

McNamara, J., Dall, S.R.X., Hammerstein, P., Leimar, O. (2016). Detection versus selection: Integration of genetic, epigenetic and environmental cues in fluctuating environments. Ecology Letters,19, 1267-1276.

Leimar, O., Dall, S.R.X., Hammerstein, P., McNamara, J.M. (2016). Genes as cues of relatedness and social evolution in heterogeneous environments. PLoS Computational Biology, 12(6): e1005006.

Zug, R. & Hammerstein, P. (2015). Wolbachia and the insect immune system: what reactive oxygen species can tell us about the mechanisms of Wolbachia-host interactions. Frontiers in Microbiology, 6, 1201.

Bossan, B., Jann, O. & Hammerstein, P. (2015). The evolution of social learning and its economic consequences. Journal of Economic Behavior & Organization, 112, 266-288.

Zug, R. & Hammerstein, P. (2015). Bad guys turned nice? A critical assessment of Wolbachia mutualisms in arthropod hosts. Biological Reviews, 90, 89-111.

Hammerstein, P. & Leimar, O. (2014). Evolutionary game theory in biology. In Handbook of Game Theory with Economic Applications, Vol. 4, eds. H.P. Young & S. Zamir, pp. 575-617. Amsterdam, Elsevier.

Telschow, A., Hilgenboecker, K. & Hammerstein, P., Werren, J.H. (2014). Dobzhansky-Muller and Wolbachia-induced incompatibilities in a diploid genetic system. PLoS ONE, 9(4), e95488.

Werner, G.D.A., Strassmann, J.E., Ivens, A.B.F., Engelmoer, D.J.P., Verbruggen, E., Queller, D.C., Noë, R., Johnson, N.C., Hammerstein, P. Kiers, E.T. (2014). The evolution of microbial markets. Proceedings of the National Academy of Sciences, 111, 1237-1244.

Bossan, B., Hammerstein, P. & Koehncke, A. (2013). We were all young once: an intragenomic perspective on parent-offspring conflict. Proceedings of the Royal Society B., 280, 1471-2954.

Zug, R., Koehncke, A. & Hammerstein, P. (2012). Epidemiology in evolutionary time: The case of Wolbachia horizontal transmission between arthropod host species. Journal of Evolutionary Biology, 25, 2149-2160.

Hammerstein, P. (2012). Towards a Darwinian theory of decision making: Games and the biological roots of behavior. In Evolution and Rationality, eds. K. Binmore & S. Okasha, pp. 7-22. Cambridge, NY: Cambridge University Press

Zug, R. & Hammerstein, P. (2012). Still a host of hosts for Wolbachia: Analysis of recent data suggests that 40% of terrestrial arthropod species are infected. PLoS ONE, 7(6), e38544.

Hammerstein, P. & Stevens, J.R. (2012). Six reasons for invoking evolution in decision theory. In Evolution and the Mechanisms of Decision Making, eds. P. Hammerstein & J.R. Stevens, in press. Cambridge, MA: MIT Press.

Hammerstein, P. & Boyd, R. (2012). Learning, cognitive limitations, and the modeling of social behavior. In Evolution and the Mechanisms of Decision Making, eds. P. Hammerstein & J.R. Stevens, pp. 319-343. Cambridge, MA: MIT Press.

Flack, J.C., Hammerstein, P. & Krakauer, D.C. (2012). Robustness in Biological and Social Systems. In Evolution and the Mechanisms of Decision Making, eds. P. Hammerstein & J.R. Stevens, in press. Cambridge, MA: MIT Press.

Bossan, B., Koehncke, A. & Hammerstein, P. (2011). A new model and method for understanding Wolbachia-induced cytoplasmic incompatibility. PLoS ONE, 6(5), e19757.

Klotz, C., Ziegler, T., Figueiredo, A.S., Rausch, S., Hepworth, M.R., Obsivac, N., Sers, C., Lang, R., Hammerstein, P., Lucius, R. & Hartmann, S. (2011). A helminth immunomodulator exploits host signaling events to regulate cytokine production in macrophages. PLoS Pathogens, 1(2), e1001248.

Leimar, O. & Hammerstein, P. (2010). Cooperation for direct fitness benefits. Philosophical Transactions of the Royal Society B, 365, 2619-2626.

Figueiredo, A.S., Höfer, T., Klotz, C., Sers, C., Hartmann, S., Lucius, R. & Hammerstein, P. (2009). Modelling and simulating IL-10 production and regulation by macrophages after stimulation with an immunomodulator of parasitic nematodes. FEBS Journal, 276, 3454-3469.

Hagen, E.H. & Hammerstein, P. (2009). Did Neanderthals and other early humans sing? Insights from the territorial advertisements of primates, lions, hyenas, and wolves. Musicae Scientiae (special issue: music and evolution). 291-320.

Hagen, E.H., Sullivan, R.J., Schmidt, R., Morris, G., Kempter, R. & Hammerstein, P. (2009). Ecology and neurobiology of toxin avoidance and the paradox of drug reward. Neuroscience, 160, 69-84.

Koehncke, A., Telschow, A., Werren, J.H. & Hammerstein, P. (2009). Life and death of an influential passenger: Wolbachia and the evolution of CI-modifiers by their hosts. PLoS ONE 4(2), e4425.

Hagen, E.H., Watson, P.J. & Hammerstein, P. (2008). Gestures of despair and hope: A view on deliberate self-harm from economics and evolutionary biology. Biological Theory, 3, 123–138.

Kobayashi, Y., Hammerstein, P. & Telschow, A. (2008). The neutral effective migration rate in a mainland-island context. Theoretical Population Biology, 74, 84-92.

Hilgenboecker, K., Hammerstein, P., Schlattmann, P., Telschow, A. & Werren, J.H. (2008). How many species are infected with Wolbachia? - a statistical analysis of current data. FEMS Microbiology Letters, 281, 215–220.

Sullivan, R.J., Hagen, E.H. & Hammerstein, P. (2008). Revealing the paradox of drug reward in human evolution. Proceedings of the Royal Society B, 275, 1231–1241.

Engelstädter, J., Hammerstein, P. & Hurst, G.D.D. (2007). The evolution of endosymbiont density in doubly infected hosts. Journal of Evolutionary Biology, 20, 685-695.

Hammerstein, P. (2007). Interview on game theory. In Game theory: 5 questions. Hendricks, V.F. & Guldborg Hansen, P. eds., pp. 75 – 95, Automatic Press.

Telschow, A., Flor, M., Kobayashi, Y., Hammerstein, P. & Werren, J.H. (2007). Wolbachia-induced unidirectional CI and speciation: Mainland-island model. PLoS ONE, 2(8), e701.

Flor, M., Hammerstein, P. & Telschow, A. (2007). Wolbachia-induced unidirectional cytoplasmic incompatibility and the stability of infection polymorphism in parapatric host populations. Journal of Evolutionary Biology, 20, 696-706.

Telschow, A., Flor, M., Kobayashi, Y., Hammerstein, P. & Werren, J.H. (2007). Wolbachia-induced unidirectional CI and speciation: Mainland-island model. PLoS ONE, 2(8), e701.

Hammerstein, P. & Leimar, O. (2006). Cooperating for direct fitness benefits. Journal of Evolutionary Biology, 19, 1400-1402

Hammerstein, P., Laubichler, M. & Hagen, E.H. (2006). The strategic view of biological agents. Biological Theory, 1, 191-194.

Telschow, A., Engelstädter, J., Yamamura, N., Hammerstein, P. & Hurst, G.D.D. (2006). Asymmetric gene flow and constraints on adaptation caused by sex ratio distorters. Journal of Evolutionary Biology, 19, 869-878.

Hammerstein, P., Hagen, E.H., Herz, A.V.M. & Herzel, H. (2006). Robustness: A key to evolutionary design. Biological Theory, 1, 90-93.

Leimar, O., Hammerstein, P. & Van Dooren, T.J.M. (2006). A new perspective on developmental plasticity and the principles of adaptive morph determination. American Naturalist, 167, 367-376.

Hagen, E.H. & Hammerstein, P. (2006). Game theory and human evolution: A critique of some recent interpretations of experimental games. Theoretical Population Biology, 69, 339-348.

Hammerstein, P. & Hagen, E.H. (2005). The second wave of evolutionary economics in biology. Trends in Ecology and Evolution, 20, 604-609.

Kowald, A., Jendrach, M., Pohl, S., Bereiter-Hahn, J. & Hammerstein, P. (2005). On the relevance of mitochondrial fusions for the accumulation of mitochondrial deletion mutants: A modelling study. Aging Cell, 4, 273-283.

Jendrach, M., Pohl, S., Vöth, M., Kowald, A., Hammerstein, P. & Bereiter-Hahn, J. (2005). Morpho-dynamic changes of mitochondria during ageing of human endothelial cells. Mechanisms of Ageing and Development, 126, 813-821.

Telschow, A., Hammerstein, P. & Werren, J.H. (2005). The effect of Wolbachia versus genetic incompatibilities on reinforcement and speciation. Evolution, 59, 1607-1619.

Hammerstein, P. (2005). Strategic analysis in evolutionary genetics and the theory of games. Journal of Genetics, 84, 7-12.

Leimar,O., van Dooren, T. J.M. & Hammerstein, P. (2004). Adaptation and constraint in the evolution of environmental sex determination, Journal of Theoretical Biology, 227, 561-570.

Engelstädter, J., Telschow, A. & Hammerstein, P. (2004). Infection dynamics of different Wolbachia-types within one host population. Journal of Theoretical Biology, 231, 345-355.

Hammerstein, P. (2003). Why is reciprocity so rare in social animals? A protestant appeal. In Genetic and Cultural Evolution of Cooperation, ed. P. Hammerstein, pp. 84-93, Cambridge, MA: MIT Press.

Bowles, S. & Hammerstein, P. (2003). Does market theory apply to biology? In Genetic and Cultural Evolution of Cooperation, ed. P. Hammerstein, pp. 153-165, Cambridge, MA: MIT Press.

Telschow, A., Hammerstein, P. & Werren, J.H. (2002). The effect of Wolbachia on genetic divergence between populations: Models with two-way migration. American Naturalist, 160, S54-S66.

Leimar, O. & Hammerstein, P. (2001). Evolution of cooperation through indirect reciprocity. Proceedings of the Royal Society, B, 268, 745-753.

Hammerstein, P. (2001). Economic behaviour in humans and other animals. In Economics in Nature, eds. R. Noë, J.A.R.A.M. van Hooff & P. Hammerstein, 1-19. Cambridge: Cambridge University Press.

Hammerstein, P. (2001). Evolutionary adaptation and the economic concept of bounded rationality - a dialogue. In Bounded Rationality: The Adaptive Toolbox, eds. G. Gigerenzer & R. Selten, pp. 71-81. MIT Press.

Hammerstein, P. (1998). What is evolutionary game theory? In Game Theory and Animal Behavior, eds. L.A. Dugatkin & H.K. Reeve, pp. 3- 15. New York: Oxford University Press.

Hammerstein, P. (1996). Streetcar theory and long-term evolution. Science, 273, 1032.

Hammerstein, P. (1996). Darwinian adaptation, population genetics and the streetcar theory of evolution. Journal of Mathematical Biology, 34, 511-532.

Kuhn, H.W., Harsanyi, J.C., Selten, R., Weibull, J.W., van Damme, E., Nash, J.F. & Hammerstein, P. (1995). The work of John F. Nash Jr. in game theory: Nobel Seminar, December 8, 1994. Duke Journal of Mathematics, 81, 1-29. Also published 1996 in the Journal of Economic Theory, 69, 153-185.

Noë, R. & Hammerstein, P. (1995). Biological markets. Trends in Ecology and Evolution, 10, 336-339.

Noë, R. & Hammerstein, P. (1994). Biological markets: Supply and demand determine the effect of partner choice in cooperation, mutualism and mating. Behavioural Ecology and Sociobiology, 35, 1-11.

Hammerstein, P. & Selten, R. (1994). Game theory and evolutionary biology. In Handbook of Game Theory with Economic Applications, Vol.2, eds. R.J. Aumann & S. Hart, pp. 929-993. Amsterdam: Elsevier.

Friedman, J.E. & Hammerstein, P. (1991). To trade, or not to trade; that is the question. In Game Equilibrium Models I: Evolution and Game Dynamics, ed. R. Selten, pp. 257-275. Berlin: Springer Verlag.

Hammerstein, P. & Riechert, S.E. (1988). Payoffs and strategies in territorial contests: ESS analyses of two ecotypes of the spider Agelenopsis aperta. Evolutionary Ecology, 2, 115-138.

Hammerstein, P. & Parker, G.A. (1987). Sexual selection: Games between the sexes. In Sexual Selection: Testing the Alternatives, eds. J.W. Bradbury & M.B. Andersson, pp. 119-142. Dahlem Konferenzen. Chichester: John Wiley & Sons.

Parker, G.A. & Hammerstein, P. (1985). Game theory and animal behaviour. In Evolution - Essays in Honour of John Maynard Smith, eds. P.J. Greenwood, P.H. Harvey & M. Slatkin, pp. 73-94. Cambridge: Cambridge University Press.

Hammerstein, P. (1984). The biological counterpart to non-cooperative game theory. Nieuw Archief voor Wiskunde (4), 2, 137-149.

Selten, R. & Hammerstein, P. (1984). Gaps in Harley's argument on evolutionarily stable learning rules and in the logic of “tit for tat”. The Behavioral and Brain Sciences, 7, 115-116.

Riechert, S.E. & Hammerstein, P. (1983). Game theory in the ecological context. Annual Review of Ecology and Systematics, 14, 377-409.

Hammerstein, P. & Parker, G.A. (1982). The asymmetric war of attrition. Journal of Theoretical Biology, 96, 647-682.

Hammerstein, P. (1981). The role of asymmetries in animal contests. Animal Behaviour, 29, 193-205.

Biographical sketch