Energetics of collective movement in vertebrates.

The collective directional movement of animals occurs over both short distances and longer migrations, and is a critical aspect of feeding, reproduction and the ecology of many species. Despite the implications of collective motion for lifetime fitness, we know remarkably little about its energetics. It is commonly thought that collective animal motion saves energy: moving alone against fluid flow is expected to be more energetically expensive than moving in a group. Energetic conservation resulting from collective movement is most often inferred from kinematic metrics or from computational models. However, the direct measurement of total metabolic energy savings during collective motion compared with solitary movement over a range of speeds has yet to be documented. In particular, longer duration and higher speed collective motion must involve both aerobic and non-aerobic (high-energy phosphate stores and substrate-level phosphorylation) metabolic energy contributions, and yet no study to date has quantified both types of metabolic contribution in comparison to locomotion by solitary individuals. There are multiple challenging questions regarding the energetics of collective motion in aquatic, aerial and terrestrial environments that remain to be answered. We focus on aquatic locomotion as a model system to demonstrate that understanding the energetics and total cost of collective movement requires the integration of biomechanics, fluid dynamics and bioenergetics to unveil the hydrodynamic and physiological phenomena involved and their underlying mechanisms.