Context-dependent variability in the components of fish escape response: integrating locomotor performance and behavior.

Escape responses are used by most fish species in order to avoid predation. Escape responses include a number of behavioral and kinematic components, such as responsiveness, reaction distance, escape latency, directionality, and distance-derived performance. All of these components can contribute to escape success. Work on the context-dependent variability has focused on reaction distance, and suggests that this component is largely determined by the relative cost and benefits of escaping (economic hypothesis). For example, reaction distance was found to depend on many factors related to perceived risk and cost of escaping, such as the attack speed and size of the predators, the proximity to refuges, and engagement in other activities (e.g., feeding). Evidence from many behavioral, kinematic, and physiological studies suggest that performance in other components of the escape response is also not always maximized. For example, escape latencies may increase in the presence of schooling neighbors, and escape speed is higher in fish that have been subject to higher predation pressure. In addition, all escape components are further modulated by the effect of environmental factors. Variability in escape components can be interpreted by using both ultimate and proximate explanations, for example, the effect of stimulus strength on escape latency can be interpreted as the triggering neural threshold varying with stimulus strength (proximate explanation) and high intensity stimuli representing higher risk to the prey (ultimate explanation). An integrative approach is suggested for a full, ecologically relevant, assessment of escape performance in fish.