This chapter provides an overview of studies on the causal analysis of behavior systems in the 40 years since Tinbergen (1963) published his views on the aims and methods of ethology. I begin with some comments on Tinbergen's conception of causation. It is then noted that, while causal work investigating the neural, hormonal, and genetic bases of behavior is flourishing, work being conducted at a strictly behavioral level of analysis has declined greatly in the past 40 years. Nonetheless, most recent research on animal cognition and applied ethology is still being carried out at a behavioral level of analysis and examples of both types of research are presented: memory mechanisms of food-storing birds and decisions of spider-eating jumping spiders as well as feather pecking in fowl and animal welfare issues are all briefly discussed. Finally, I discuss the similarities between neural network modeling and early ethological models of motivation, and then show how a modern version of Lorenz's model of motivation can account for current research findings on dustbathing in chickens and sleep in humans. I conclude that valuable information can still be obtained by research at a behavioral level of analysis.

CONCEPTION OF CAUSATION

Tinbergen uses the word “causation” to refer exclusively to what many behavioral biologists currently call “proximate causation”: the immediate effects that external and internal factors have on the occurrence of behavior. He begins his analysis of causation by referring to three statements made by Lorenz (1937), which he then proceeds to discuss.