Costs of locomotion and vertic dynamics of cephalopods and fish

The world's oceans are three-dimensional habitats that support high diversity and biomass. Because the densities of most of the constituents of life are greater than that of seawater, planktonic and pelagic organisms had to evolve a host of mechanisms to occupy the third dimension. Some microscopic organisms survive at the surface by dividing rapidly in vertically well mixed zones, but most organisms, small and large, have antisinking strategies and structures that maintain vertical position and mobility. All of these mechanisms have energetic costs, ranging from the "foregone metabolic benefits" and increased drag of storing high-energy, low-density lipids to direct energy consumption for dynamic lift. Defining the niches in the mesopelagic zone, understanding evolution, and applying such ecological concepts as optimal foraging require good estimates of these costs. The extreme cases above are reasonably well quantified in fishes, but the energetic costs of dynamic physiological mechanisms like swim bladders are not; nor are the costs of maintaining vertical position for the chief invertebrate competitors, the cephalopods. This article evaluates a matrix of buoyancy mechanisms in different circumstances, including vacuum systems and ammonium storage, based on new data on the metabolic cost of creating buoyancy in Sepia officinalis.