Avian emotions: Comparative perspectives on fear and frustration

Emotions are complex reactions that allow individuals to cope with significant positive and negative events. Research on emotion was pioneered by Darwin's work on emotional expressions in humans and animals. But Darwin was concerned mainly with facial and bodily expressions of significance for humans, citing mainly examples from mammals (e.g., apes, dogs, and cats). In birds, emotional expressions are less evident for a human observer, so a different approach is needed. Understanding avian emotions will provide key evolutionary information on the evolution of related behaviors and brain circuitry. Birds and mammals are thought to have evolved from different groups of Mesozoic reptiles, theropod dinosaurs and therapsids, respectively, and therefore, their common ancestor is likely to be a basal reptile living about 300 million years ago, during the Carboniferous or Permian period. Yet, birds and mammals exhibit extensive convergence in terms of relative brain size, high levels of activity, sleep/wakefulness cycles, endothermy, and social behavior, among others. This article focuses on two basic emotions with negative valence: fear and frustration. Fear is related to the anticipation of dangerous or threatening stimuli (e.g., predators or aggressive conspecifics). Frustration is related to unexpected reward omissions or devaluations (e.g., loss of food or sexual resources). These results have implications for an understanding of the conditions that promote fear and frustration and for the evolution of supporting brain circuitry.