The rates of metabolism in animals vary tremendously throughout the biosphere. The origins of this variation are a matter of active debate with some scientists highlighting the importance of anatomical or environmental constraints, while others emphasize the diversity of ecological roles that organisms play and the associated energy demands. Here, we analyse metabolic rates in diverse marine taxa, with special emphasis on patterns of metabolic rate across a depth gradient, in an effort to understand the extent and underlying causes of variation. The conclusion from this analysis is that low rates of metabolism, in the deep sea and elsewhere, do not result from resource (e.g. food or oxygen) limitation or from temperature or pressure constraint. While metabolic rates do decline strongly with depth in several important animal groups, for others metabolism in abyssal species proceeds as fast as in ecologically similar shallow-water species at equivalent temperatures. Rather, high metabolic demand follows strong selection for locomotory capacity among visual predators inhabiting well-lit oceanic waters. Relaxation of this selection where visual predation is limited provides an opportunity for reduced energy expenditure. Large-scale metabolic variation in the ocean results from interspecific differences in ecological energy demand.Keywords: metabolism; deep sea; scaling; oxygen consumption; locomotion; marine
INTRODUCTIONThe rates of metabolic processes in animals vary tremendously throughout the biosphere (e.g. Bennett 1991; Seibel 2007). The origins and scope of this variation are a matter of active debate. Some emphasize geometric and environmental constraints or resource limitation as its source (e.g. Gillooly et al. 2001;Brown et al. 2004), while others emphasize the diversity of ecological roles that organisms play and the associated energy demands (Childress 1995;Suarez 1996;Reinhold 1999;Clarke 2004;Seibel 2007) as a primary driver of metabolic variation. With this in mind, the present paper addresses three seemingly simple questions: (i) what is the extent of variation in the rate of metabolism across diverse taxa and environments, (ii) what environmental and ecological demands act on organisms to drive selection for high metabolic capacity, and (iii) under what environmental circumstances might such selection be relaxed or capacity be constrained?These questions have been pursued vigorously for decades but the clarity of the debate has been diminished by the subtlety of differences in capacity between the taxa most thoroughly studied, that often