‘Animal Behavioral Economics’: Lessons Learnt From Primate Research
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The paper gives an overview of primate research and the economic-ethical lessons we can derive from it. In particular, it examines the complex, multi-faceted, and partially conflicting nature of (non-)human primates. Our closest living relatives, the chimpanzees and bonobos, are apparently walking on two legs: a selfish and a groupish leg. Given evolutionary continuity and gradualism between monkeys, apes and humans, human primates seem to be also bipolar apes. They, too, tend to display a dual structure: there seems to be a pro-social and a self-interested side to our species; a bipolar tension seems to exist between competition and cooperation respectively between self-interest and the common good. We are apparently at the same time Homines oeconomici and Homines culturali. Our inner ape tries to combine self-interested and common good motives. Based on de Waal’s Russian doll model, the essay investigates the evolutionary origins of morality and eusociality. With the help of selected case studies stemming from behavioral sciences/economics, the paper illustrates examples of empathy, altruism, reciprocal fairness, pro-social and other-regarding preferences, inequity aversion, and altruistic punishment in (non-)human primates. Beside this selfless and groupish side, the paper also reflects on the self-interest and egoistic nature of (non-)human primates and the behavioral and cognitive differences between monkeys, apes and humans.
Manuel Worsdorfer’s “Animal Behavioral Economics” is about the findings from non-human primate research. Somewhat surprisingly, though, it has much to say about humans and behavioral economics, particularly humans’ core motivations and ethical behavior. Therefore, it is very much of interest to social scientists, especially economists.
Worsdorfer has provided a very competent overview of non-human primate (NHP) research that focuses on the behaviors and motivations of a number of types of monkeys and apes and the ethical lessons that can be drawn from these. Humans and NHPs have much in common insofar as they have a bipolar aspect. That is, they pursue their self-interests (rational utility maximizing and competitive behavior), and they also pursue other-regarding, prosocial behavior oriented toward the common good. Nevertheless, humans are different from NHPs. Many of the differences reflect the fact that human primates have evolved a few steps beyond NHPs, and therefore, have in important respects greater social and psychological capacities.
Worsdorfer provides a good overview of the research on empathy, theory of mind, and mirror neurons related to NHPs and other animals in comparison to each other and to humans. By virtue of natural selection, all primates have brains with some degree of empathetic capacity. Of all the great apes, bonobos have the highest level of empathy. NHPs are in many ways “in tune with their fellow beings” with regard to feeling their distress, pleasure, pain, delight, etc. NHPs also have an ability to understand others’ goals, intentions, and perceptions, and are able to help others in a targeted, insightful way. Nevertheless, Worsdorfer concludes that “humans grasp how others feel and what they might need more fully than any other animal species.” Despite the fact that “humans have more of everything,” Worsdorfer observes that “there are no sharp dividing lines … between humans and apes.” The differences between humans and NHPs reflect the “biological gradualism” of evolution. Humans no doubt are more advanced than their primate cousins in regard to trust, reputation building, moral development, deliberation, and reasoning.
The research findings have important implications for morality. Worsdorfer observes that “moral behavior is a result of the ‘compassionate instincts’” that are deeply ingrained in the biological heritage of primates. Morality is “not uniquely human; it was not purposely or consciously developed.” Primate morality is a product of evolutionary processes that “favor those animals that … help each other, form trusting alliances, and establish and maintain coalitionary support.”
Interestingly, the NHP research findings are roughly consistent with dual motive theory and the model of the triune brain. In this model, the brain consists of three modules, the evolutionary oldest and lowest part is the reptilian brain, then comes the paleo-mammalian brain, followed by the neo-mammalian brain on top. According to dual motive theory, the neo-mammalian brain serves to balance self-interest motivation deriving from the reptilian brain with the other-interested motivation stemming from the old mammalian brain. Of the primates, humans have the largest and most well developed neo-mammalian (neocortex) brain, and thus would be expected to have a better integration of the two motives and better brain functioning overall.
Worsdorfer summarizes many of the NHP research findings, some of which utilize experiments in the form of the ultimatum and dictator games; other findings come from observations of NHPs in the field. Each of the experiments and observations provide useful perspectives that help to understand further the mental capabilities of the different NHPs and how they differ (or not) from human capabilities. As a behavioral economist, this article was very useful in its view of humans as primates, all of whom share the same fundamental motivations and biology. The perspectives of this paper can help economists gain more realistic appreciation of the capabilities and motivations of the human actor in economics.
John F. Tomer
Emeritus Professor of Economics
As a non-expert in primate research and zoology, I learned a lot from reading this very rich and interesting paper. The depth and scope of Worsdörfer’s analysis is impressive, and the links drawn between human and animal behaviour are plausible and relevant for economics and more generally for the social sciences. I have one brief observation to make and three detailed comments.
My general observation is that it is not obvious that ‘ethical’ lessons can be learned from the research under discussion. Primate research tells us a lot about the nature and origins of pro-social behaviour in animals. Paired with the evolutionary gradualism described by Worsdörfer, this research provides a potentially important resource for anyone wishing to combat the idea of homo-economicus as a description of human nature. In this sense, the paper reinforces the critique of mainstream economic theory insofar as it espouses a vision of homo-economicus as a positive theory of human behaviour. However the many variants of rational choice theory that currently exist across the social sciences are often presented as normative theories (not theories of human nature, but theories of what human beings should do under idealised circumstances). It is not clear to me how the zoological evidence presented in the paper can be used to challenge these normative positions. More generally, I don’t see how animal research can tell us very much about what is or isn’t morally right and thus, it seems a misnomer to speak about ‘ethical lessons’.
Three specific additional comments:
1. Page 2 (top): the definition of MI presented is a little unusual in particular the reference to disinterested and unemotional choices, and the idea of representative individuals.
2. Page 2 (middle): it seems very strange to refer to the idea of homo culturalis in heterodox economics and then cite Akerlof & Kranton (2010) when there is a very big heterodox literature on the topic (see for example Davis 2003).
3. Section 3.1 is somewhat repetitive and could easily be shortened.
Davis, J. B. (2003). The theory of the individual in economics: Identity and value. Routledge.