The synthetic approach to the study of spatial memory: have we properly addressed Tinbergen's "four questions"?

In 1963, Niko Tinbergen suggested that to truly understand the behavior of an animal, the ultimate causes (e.g., adaptive value, evolutionary history) as well as the proximate mechanisms (e.g., neurobiology, development) that result in the production of the behavior must be understood in an integrated framework. We examine whether the study of spatial memory in food storing birds has adequately addressed Tinbergen's questions and highlight the work of Sara Shettleworth, who has made a tremendous contribution to this area of study, and whom this issue honors. Our conclusion is that while the study of food caching and spatial memory in birds has been a very good model of a program of research that has addressed Tinbergen's questions, additional work remains.