Optimal diving behaviour and respiratory gas exchange in birds

This review discusses the advancements in our understanding of the physiology and behaviour of avian diving that have been underpinned by optimal foraging theory and the testing of optimal models. To maximise their foraging efficiency during foraging periods, diving birds must balance numerous factors that are directly or indirectly related to the replenishment of the oxygen stores and the removal of excess carbon dioxide. These include (1) the time spent underwater (which diminishes the oxygen supply, increases carbon dioxide levels and may even include a build up of lactate due to anaerobic metabolism), (2) the time spent at the surface recovering from the previous dive and preparing for the next (including reloading their oxygen supply, decreasing their carbon dioxide levels and possibly also metabolising lactate) and (3) the trade-off between maximising oxygen reserves for consumption underwater by taking in more air to the respiratory system, and minimising the energy costs of positive buoyancy caused by this air, to maximise the time available underwater to forage. Due to its importance in avian diving, replenishment of the oxygen stores has become integral to models of optimal diving, which predict the time budgeting of animals foraging underwater. While many of these models have been examined qualitatively, such tests of predictive trends appear fallible and only quantifiable support affords strong evidence of their predictive value. This review describes how the quantification of certain optimal diving models, using tufted ducks, indeed demonstrates some predictive success. This suggests that replenishment of the oxygen stores and removal of excess carbon dioxide have significant influences on the duration of the surface period between dives. Nevertheless, present models are too simplistic to be robust predictors of diving behaviour for individual animals and it is proposed that they require refinement through the incorporation of other variables that also influence diving behaviour such as, perhaps, prey density and predator avoidance.