3.8 Tinbergen’s Four Questions
Nikolaas “Niko” Tinbergen (1907-1988) was a Dutch biologist and is considered one of the founders of modern ethology. In 1963, his seminal paper titled “On aims and methods of Ethology” sought to build a connection between the study of biology and the study of behavior. The resulting framework is known as Tinbergen’s Four Questions.
Tinbergen formalized the field of ethology by offering a multi-level explanation to address “how” and “why” behaviors evolve and persist (ultimate causation) and how they arise in an individual (proximate causation) (Tinbergen, 1963; Bateson & Laland, 2013). These four questions can be distilled as follows:
- Proximate Causation (regarding individual organisms):
- Ultimate Causation (regarding species level traits):
- Phylogeny: What is the evolution of the behavior? Is the behavior derived (new to this species) or ancestral (present in immediate, phylogenetic ancestors)?
- Adaptive Value: What is the survival function of the behavior? What is it about the behavior that confers a reproductive advantage over others?
The mechanism of the behavior can be explained as the biology that gives rise to or results from the behavior’s expression. For example, in anxiety this may be the racing heart or sweaty palms. It may also be the adrenaline pumping into your body. Ontogeny speaks to the developmental history, for example how learning or culture shapes whether you embrace anxiety as a signal about your environment or try to suppress it. The adaptive value speaks to whether the behavior conferred a survival or reproductive advantage to previous generations of the species. In truth, anxiety is deeply connected to the fight or flight system – and it is always good to flee when a predator is chasing you. Phylogeny is the evolutionary past of a behavior within the species and those to whom the species is most immediately related. For example, all mammals nurse their young (ancestral), but only humans will use tools like bottles to deliver milk (derived).
Proximate Questions |
Ontogeny(Development/Culture) What learning or developmental history shapes the way this behavior is expressed by an individual organism? |
Mechanism(Biology) What biological triggers or changes bring about this behavior in an individual organism? |
Ultimate Questions |
Phylogeny(Evolutionary History) Is this behavior new to the target species or is it shared with others who share a common ancestry? |
Adaptive Value(Function/Survival Purpose) Why does this behavior persist in the species? How did it help the species succeed? |
An integrative use of Tinbergen’s four questions becomes crucial to a holistic understanding of the development of any trait. Often, we can ask a question as “why” and Tinbergen gives us four potential reasons. Let’s practice this now.
Watch the following video for a practical explanation of Tinbergen’s Four Questions, specifically as they apply to bird song.
the study of human behavior and social organization from a biological perspective.
causes that are the mechanisms directly underlying the behavior in an individual organism.
developmental influences on behavior; include cultural influences and learning history, and may include aspects of physiological growth.
biological triggers or results of individual behavior occurring in response to environmental or internal stimuli.
refers to the behavior's evolutionary significance or phylogeny; how the behavior enhances reproductive fitness in the species.
the branch of biology that deals with classifying the evolutionary development and diversification of a species or group of organisms.
a trait that is new to the species being studied; a trait that is not present in any of the immediate ancestors of the species.
a trait that was present in the immediate ancestors of a species; may be evolutionarily ancient or within recent ancestral species.
the reason a trait exists within a species; the reproductive or survival benefit it confers.