Niko Tinbergen, a pioneer in the study of animal behavior, wisely observed that four questions need to be asked for all products of evolution1.

  1. Given that a trait is an adaptation, what is its function that contributes to survival and reproduction?
  2. Given that evolution is a historical process, what is the phylogeny of the trait?
  3. Given that all traits (including behaviors) have a physical basis, what is the mechanism of the trait?
  4. Given that all traits must come into being during the lifetime of the organism, what is the development of the trait?

Tinbergen’s four questions apply to any variation-and-selection process, including but not restricted to genetic evolution2.  Accordingly, they can be insightful for the study of moral universals and particulars as products of human genetic and cultural evolution.

Function: The most general statement that can be made about human morality is a functional one: In virtually all cultures, most people have a sense of right and wrong that corresponds to the welfare of their groups. Also, most people create, abide by, and enforce norms on the basis of what they regard as right and wrong. Notice that this generality is statistical in nature. It admits the possibility that some individuals might not qualify as moral. For example, psychopaths are said to lack a sense of right and wrong, treating everything as instrumental to their desires3. In most experimental games that measure cooperative behavior, a sizable fraction of individuals don’t cooperate and/or don’t punish norm transgressions4. Nevertheless, enough individuals behave morally in virtually all cultures so that the cultures function as moral systems.

Tinbergen’s four questions apply to any variation-and-selection process, including but not restricted to genetic evolution. Accordingly, they can be insightful for the study of moral universals and particulars as products of human genetic and cultural evolution.

Phylogeny: The reason that we are psychologically endowed to behave morally, to the extent that we do, is because of a historical process of between-group selection. As Darwin conjectured long ago, individuals who behave morally are vulnerable to more self-serving individuals within their own groups, but groups of individuals who behave morally robustly out-compete groups whose members can’t pull together. The fact that between-group selection (favoring the traits associated with morality) is often opposed by within-group selection (favoring the traits associated with immorality) explains why all of us behave immorally at least some of the time and some of us more than others. Insofar as different environments call for different behaviors to benefit a given group, the specific behaviors that count as moral can be highly variable. Also, not everything that evolves is an adaptation. There are byproducts, products of drift, and mismatches (adaptive in past but not present environments) for cultural in addition to genetic evolution. Thus, Tinbergen’s Phylogeny question can explain a lot of moral particularism.

Mechanism: What takes place in our brains when we behave morally? The answer might be “it depends”. One person might behave out of a sense of duty. Another might take pleasure in helping others. Another might be trying to earn a ticket to heaven. There is inherently a one-to-many relationship between the function of a trait and the proximate mechanisms that evolve to cause it. This is important because philosophers often reason on the basis of their own moral intuition as if it must be culturally universal. There is no warrant for this assumption from an evolutionary perspective. We must realize that the proverb “there are many ways to skin a cat” applies to the mechanisms underlying moral behaviors along with many other kinds of behaviors.

Development: Our core psychological ability to function as moral agents might qualify as universal or nearly so, with developmental stages that are correspondingly universal. However, the particular moral systems and their underlying mechanisms that evolve in any particular culture will also have particularistic developmental pathways. This will require a rethinking of some stage theories of human moral development. For example, in Kohlberg’s theory of moral development5, the highest stage is driven by universal ethical principles. When this “stage” is reconceptualized as a moral system that competes against other moral systems, it requires very special conditions to evolve, which accounts for the fact that most individuals and cultures don’t achieve it. Creating such a moral system is an important normative goal that I share, but there is no warrant for calling it a stage in a developmental sense.

While the topic deserves much more than a short commentary, Tinbergen’s four questions might prove as useful for organizing the study of morality as for all other products of evolution.


  1. Tinbergen, N. (1963). On aims and methods of ethology. Zeitschrift Für Tierpsychologie, 20, 410–433.
  2. A lecture that I frequently give on Tinbergen’s four questions is available online at https://www.youtube.com/watch?v=yYrSQ3IULhk
  3. O’Conner, L. E., Berry, J. W., Lewis, T. B., & Stiver, D. J. (2011). Empathy-based pathogenic guilt, pathological altruism, and psychopathology. In B. Oakley, A. Knafo, G. Madhavan, & D. S. Wilson (Eds.), Pathological Altruism (pp. 10–30). Oxford: Oxford University Press.
  4. Fehr, E., Fischbacher, U., & Gachter, S. (2002). Strong reciprocity,human cooperation, and the enforcement of human social norms. Human Nature, 415, 137–140.
  5. Kohlberg, L. (1984). Essays on Moral Development: Vol. 2. The Psychology of Moral Development. San Francisco: Harper and Row.

This article is from TVOL’s project titled “This View of Morality: Can an Evolutionary Perspective Reveal a Universal Morality?” You can download a PDF of the project [here], comment on this article below, or comment on the project as a whole in the Summary and Overview.

Published On: May 17, 2018

David Sloan Wilson

David Sloan Wilson

David Sloan Wilson is president of Prosocial World and SUNY Distinguished Professor Emeritus of Biology and Anthropology at Binghamton University. He applies evolutionary theory to all aspects of humanity in addition to the rest of life, through Prosocial World and in his own research and writing.  A complete archive of his work is available at www.David SloanWilson.world. His most recent books include his first novel, Atlas Hugged: The Autobiography of John Galt III, and a memoir, A Life Informed by Evolution.



  • Mark Sloan says:


    Have you heard of Tinbergen’s four questions being formally applied to help sort out the complexities and strangeness (due to its apparent harm) of the various forms of Female Genital Mutilation?

    My interest in FGM is: 1) if it is a ‘moral’ norm (rather than just a cultural norm), then moral theories must be able to explain its origin, function, and why its harm is tolerated and 2) how understanding those origins and function may be culturally useful.

    Perhaps application of Tinbergen’s four questions could be a useful tool in making FGM less mysterious.

    • Rhea Luana Arini says:

      Dear Mark,

      I think you might find this paper of your interest as it discusses FGM’s origins and maintenance under an evolutionary point of view and, importantly, it contains a section about practical implications of the study with the aim to implement policies to end this practice.

      The Origins and Maintenance of Female Genital Modification across Africa: Bayesian Phylogenetic Modeling of Cultural Evolution under the Influence of Selection


      • Mark Sloan says:

        Hi Rhea,
        Interesting paper. Not only is there a summary of recent data but the mathematical modeling approach looks useful.
        FGM’s correlation with polygamous societies due to competition for high value mates is consistent with my own speculations.
        Thanks for suggesting.

  • Andy Norman says:

    Tinbergen’s questions seem to be a good way to structure an account of the evolution of moral universals. I don’t dispute any substantive aspect of David’s account, for I believe he understands the issues far better than I do. I will raise a couple of questions, though, just to see where they go.

    When discussing the evolution of human reasoning capacities with Michael Tomasello, we looked at the possibility that group selection may have played a role in their evolution. Michael mentioned something he called “social selection,” a process whereby a group expels a free-rider for not pulling his weight, and the exile perishes. Michael’s idea, as I understand it, is that we might be able to explain the emergence of (some) pro-social traits with social selection rather than group selection. I don’t know enough game theory to test this supposition, but I’d be interested in hearing what David thinks of it. (Full disclosure: I have no particular allergy to group selectionist accounts.)

    My other question is merely a semantic quibble, but perhaps worthy of pondering even so: Are moral universals properly thought of as “traits” at all? Certain moral dispositions, such as sympathy for (some) others, do seem to be hard-wired into almost all of us. Our capacity for sympathy is probably distributed across some kind of bell curve, however, so even within this one purportedly universal trait, we find difference. If there is a set of moral principles that are binding on all of us however, no matter how sympathetic or unsympathetic we happen to be, this set is universal in a somewhat different sense. My inclination is to call things of the latter type true moral universals, and things like our average de facto allotment of moral sympathy “traits.” That way, we’re less likely to mix up de facto traits and de jure ideals. A small matter, and only relevant in very particular circumstances…

    • Sage Gibbons says:

      Andy, I’m not sure why one would need to draw a distinction between social selection and group selection.

      The social selection you describe is a group-level process that benefits the group’s prosociality by minimizing disruptive lower-level selection that reduces the adaptive potential of cooperation and as a result, maintains prosociality’s expression within the group such that it is a viable selective factor between groups. In other words, the well documented exclusion of bad players (or simply outsiders) is a group-functional trait that also counters group cultural drift in the same way that individual mate choice counters genetic drift. It’s hard to see how a group-beneficial process that also maintains the between-group variation necessary for group selection to occur is in some way not part of group-selection.

      Caveat: My point on cultural group drift is generally limited to cases where the number of groups are small, but that’s more of an empirical contingency than a theoretical one.

  • I love Tindbergen’s four questions. 1. “What is its function that really does the work of aiding survival?” (I’m paraphrasing) My guess is that morality regulates dominance relations. It is a way of stopping bullies and cheaters with getting away with their actions. By minimizing bullying and cheating, a space is created for good behaviour, allowing people in society to exercise free choice as long as they don’t break the rules themselves. The upshot of this is that more children survive until adulthood, and groups are larger and have greater cohesion against threats. Isn’t it fascinating how global events are highlighting the importance of morality? If a bully, let’s call him “DT”, is allowed to intimidate and threaten others with impunity, he will escalate his threats and seize greater power. Morality is the only way that the group can unify together to eliminate bullies and prevent them from re-emerging.
    Questions two and four are important, but answering either one opens a can of worms.
    I would paraphrase question three as: “How does it work?” Answer: a lot like a common pool resource or CPR. Everyone knows the rules, everyone follows the rules, and everyone monitors for cheaters and bullies. Violations are detected, and violators are punished. If people are satisfied that this is working they are less likely to commit violations themselves. One should be able to see this process happening on all human scales, from neighbourhoods to the global system.

  • Jack Sawyer says:

    “There are many waya to skin a cat,” David writes. But are any of them moral? Words are powerful. We need to use them responsibly.

  • Ramesh says:

    In my opinion, origin of morality is very simple. We all have two instincts hard wired into us, into all the animals. Survival of the self survival and survival of the species. These must be very strong instinct in any species. Any species which do not have them would have died out long time ago.

    So humanity also has these two interests hard wired into its DNA. Any selfish acts we perform, getting a education, earning a living, getting rich, having children, even committing a murder, can be traced back to the instinct of self survival.

    Any charitable, compassionate, altruistic acts we perform, can be attributed to the instinct survival of the species. We do these acts because it helps other fellow human beings, that contributes to the survival of the species.

    Morality is really not any more complicated than that. There is no need to invoke an anthropomorphic God.

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