Physiological constraints on contest behaviour

1. Contests may involve injurious fighting, other types of direct physical aggression and communication. They occur over ownership access to mates and other resources that may increase an individual's attractiveness and its chance of survival. Traits that enhance resource holding potential may be the result of sexual selection, natural selection or a combination of both. 2. Agonistic behaviours are expected to be demanding to perform and costly in terms of changes in physiological state. The ability to meet the physiological costs may determine contest outcomes and constrain the intensity of agonistic activities. 3. The energetic costs have been investigated in a broad range of taxa using a variety of techniques. They include the mobilization of energy reserves, but a key cost in several taxa appears to be limited anaerobic capacity and subsequent accumulation of lactic acid. Androgens, stress hormones and neurohormones have also been shown to constrain fighting behaviour. However, due to key differences in the endocrine systems of vertebrates and invertebrates, the effects of hormones are far less consistent across taxa than in the case of metabolites. 4. Physiological constraints on fighting may vary according to their importance relative to circumstantial costs, the time-scale over which they exert their effects, their effects on different roles and their causal links with behaviour. Incorporating these factors into theoretical studies of contest behaviour may give further insights of how the costs of fighting influence agonistic behaviour.