Tinbergen’s Four Questions

Tinbergen’s Four Questions are questions that can be asked about an organism and its evolution that help to explain its adaptations and behavior. They operate via a double dual of Static vs. Dynamic and Proximate vs. Ultimate. Static is concerned with the current time whereas Dynamic considers a time series. Proximate is concerned with how the organism behaves currently or how its behavior changes over its lifetime, and Ultimate considers why the organism and its behavior/adaptations may have evolved the way they did. Both pairs are somewhat confusedly concerned with time: both Static and Proximate are concerned with either the current time or a short lifetime, and Dynamic and Ultimate are concerned with changes in that short lifetime or over evolutionary time.

Ultimate & Static: Function or Adaptation
Ultimate & Dynamic: Phylogeny or Evolution
Proximate & Dynamic: Ontogeny or Development
Proximate & Static: Mechanism or Causation

Ultimate is also called Evolutionary, to distinguish it from a connotation of telos or purpose. Static refers to the current form of the organism, and is also called Synchronic or Single Form or Snapshot or Contemporary, etc. Dynamic refers to the historical changes of the organism, and is also called Diachronic or Sequence or Historical or Chronicle, etc.

Some compare these four questions to Aristotle’s Four Causes, see for example [1] and [2]. However, [1] seems less enthusiastic than the published paper [2]. Also, I don’t agree with either completely on the assignment; it seems to me that the Efficient and Final Causes are Dynamic, and the Material and Formal causes are Static. I believe we all agree that the Efficient and Material Causes are Proximate, and Formal and Final Causes are Ultimate. See [3] for comparison. Frankly, I am guided more by the definitions of Static and Dynamic than anything else.