Ventilation Frequency Reveals the Roles of Exchange Surface Areas in Metabolic Scaling

The surface area (SA) theory proposes that resting metabolic rate (RMR) scales with body mass, which parallels the exchange SA of organisms, and that a species with a larger scaling exponent of exchange SA has a larger scaling exponent of RMR. However, the effects of exchange SA on metabolic scaling may be eclipsed because oxygen transfer across the respiratory surface is determined not only by the exchange SA but also by ventilation. We hypothesize that the scaling of both gill surface area (GSA) and ventilation frequency (VF) positively affects the scaling of metabolic rate. In six closely related species of carp maintained under the same experimental conditions, the scaling exponents of RMR and GSA were analyzed. In the goldfish, RMR scaled with body mass by an exponent significantly lower than that of GSA but not different from the exponents of GSA in the remaining five species. The scaling exponent of RMR was positively related to those of both GSA and VF among the species. In addition, the VF-corrected metabolic scaling exponent was positively related to the scaling exponent of GSA among the species. These results suggest that variations in GSA scaling and in VF scaling among species mutually affect metabolic scaling.