Abstract: The question “Why does an animal behave as it does?” can be answered in terms of ontogeny, function, phylogeny and causation. The achievements of applied ethology relative to those four approaches are reviewed, gaps in our knowledge are identified and predictions for fruitful avenues of future research are made. Ontogenic studies have been useful in the past and it is suggested that studies of the effects of early experience on the sexual behaviour of animals used in artificial breeding schemes might pay dividends. It is proposed that functional studies should be approached cautiously. More information is required on the process of domestication in order to increase the chances of success in the trend to farm exotic species. Studies on causation are likely to continue to be the mainstay of applied ethological research. It is suggested that within this category, studies on states of suffering, motivation and cognition are urgently required to answer the most pressing questions on animal welfare.