Human beings have always questioned the origin and causes of things that happen to them. The application of this curiosity for knowledge to the field of biology has given way to ethology, among other branches of science.

One of the fathers of this science is Nikolaas Tinbergen, a zoologist who made several contributions to the study of living beings. Among them, we find what is known as Tinbergen’s 4 questions , an effort to sort out the unknowns to be answered about the biology and behaviour of any animal (including humans).

What is the function of a behavior? How does it develop, evolve and what causes it? If you want to know these answers, read on.

Background: the beginnings of biology

Aristotle already stated that “to know something scientifically is to know its causes”. He also established 4 types of causes: material, formal, efficient and final . This can be considered a precedent to Tinbergen’s questions, since he intended it to be the starting point of the investigations of any researcher who wanted to study nature.

Before Tinbergen, around 1930, Julian Huxley spoke of three major problems in biology: cause, survival value, and evolution. It was Niko Tinbergen who added the fourth: ontogeny, that is, the development of each individual from birth to death. On the other hand, Ernst Mayr in 1961 spoke of proximate and ultimate cause.

What are Tinbergen’s 4 questions?

Niko Tinbergen, considered one of the fathers of ethology, was a Dutch zoologist who was born in 1907. In 1973 he was awarded the Nobel Prize in Physiology and Medicine, together with Konrad Lorenz and Kar von Frisch, for his discoveries in relation to individual and social behaviour patterns .

Tinbergen, in his article On aims and methods of ethology of 1963, raises the existence of 4 main problems of biology, or the 4 questions of Tinbergen , which are levels of biological explanation of certain phenomena of nature.

Tinbergen poses these questions in order to understand a behavior, and they are as follows.

Cause or mechanism: What is the cause of the behavior?

It represents the proximate or structural cause. It is the internal and external stimuli that trigger the behaviour .

Here, sensory receptors play a key role by allowing the information provided by such stimuli to be perceived.

Survival value: How does such behavior contribute to the survival and reproductive success of the animal?

It represents the ultimate cause. That is, the function, adaptation or adaptive advantage of the behaviour.

Ontogeny: How does such behavior develop during the animal’s life?

It has to do with the possible changes and evolution that a pattern of behavior undergoes throughout an individual’s life.

Evolution: How has behavior evolved?

Also called phylogeny. It studies the phylogenetic history of such behavior, that is, of the precursors . Thanks to this, it can be understood that the behaviour is in such a way nowadays, and not in another way.

The levels of biological explanation

Relating Tinbergen to Mayr, we see that the proximate causes (immediate in time) would encompass mechanism and ontogeny, and the evolutionary causes (more distant or distant) would encompass survival value and phylogeny.

Thus, the former would explain the structure and mechanisms of behavior, and the latter, why organisms are the way they are.

Practical example

To illustrate Tinbergen’s questions, let’s look at an example . This is a rough guide to get an idea, but the answers will always vary from case to case.

Let’s think of a child who hits others when he’s angry. Let’s analyze the components of such behavior according to Tinbergen’s 4 questions.


This may be due to irritability, low tolerance to frustration, lack of other emotional support skills, etc.


Calling attention to yourself, venting your anger, showing your irritability so that you can be attended to.


It develops and repeats itself because it has previously shown similar behaviors and these have been reinforced at some point.


The child has seen how his siblings were reinforced by such behavior, and he reproduces it.

Implications for Science

As we have seen, we can shell and analyze the components of each animal behavior that we consider , although obviously not all behaviors will have the same function, much less the same adaptive value.

There will be behaviors that are more adaptive than others, and these will probably be the ones that will be repeated in the evolutionary chain and those that will be consolidated in a more stable way in a species .

Today, 50 years after the publication of that article, Tinbergen’s 4 questions are still considered one of the most important and valuable legacies of the author, because of the importance he attaches to his comprehensive and multifaceted view of a behaviour.

Author’s vision and conclusion

Tinbergen gave a pragmatic character to his theory, as well as a logical one, which makes his work a useful and complete tool to understand behavior. He was among the first to study the adaptive meaning of behaviours that may seem useless in the first place; for example, he studied the behaviour of laughing gulls when they remove the shells of their eggs from the nest once their chicks are born.

The author considered that grouping the problems would facilitate the understanding of behaviour , and considered it a fundamental part of ethology. Anyway, he always bet not only on integrating behaviors, but also on studying them individually, thus acquiring an analytical and global vision of the behavior or the problem at the same time.

Tinbergen’s 4 questions are apparently simple, but at the same time synthetic, since they lead us to a complete understanding of a biological or behavioural phenomenon.

Bibliographic references:

  • Donal, A. (1999). The proximate and the ultimate: past, present and future. Behavioral Processes, 189-199.
  • Bateson, P. & Laland, K. (2013). Tinbergen’s four questions: an appreciation and an update. Trends in Ecology & Evolution, 28(12), 712-718.