Tinbergen's four questions, organized: a response to Bateson and Laland

摘要

‘What people think of as the moment of discovery is really the discovery of the question’, according to a comment attributed to Jonas Salk. For biology, that moment came in 1963 when Niko Tinbergen published an essay describing Four Questions that need to be answered to explain fully a biological trait. Aristotle offered similar categories, Huxley described three of them in modern terms, and Ernst Mayr paved the way with his distinction between proximate and ultimate (evolutionary) explanations 1 and 2, but Tinbergen's Four Questions have proved most valuable for biology. The 50th anniversary of their publication provides, as Bateson and Laland note, an occasion for celebration and reflection [3]. Their article focuses on possible ways the questions could be amended in light of modern advances and debates. The implications of gene–culture evolution, levels of selection, exaptation, and epigenetics for Tinbergen's questions are certainly worth discussion, as the Bateson and Laland article illustrates. However, emphasizing such controversies and subtleties can obscure Tinbergen's accomplishment, which remains vastly underappreciated.