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The metabolic-level boundaries (MLB) hypothesis and the cell metabolism (CM) hypothesis have been proposed to explain the body mass scaling of metabolic rate. The MLB hypothesis focuses mainly on the influence of the metabolic level on the relative importance of volume and surface area constraints. The CM hypothesis focuses on the variation of cell size as the body grows. The surface area to volume ratio of individual cells may vary among species with different cell sizes, by which surface area constraints on metabolic scaling may change according to the MLB hypothesis. The present study aimed to extend the MLB and the CM hypotheses by proposing that, in addition to metabolic level, the varying cell surface area constraints among species also influence the intraspecific scaling exponents. The red blood cell area (S), and intraspecific scaling exponents for resting (b R ) and maximum metabolic rates of four species of cyprinids were assessed. The scaling exponents varied among species, but mass-specific resting metabolic rates (RMR) of each species were similar. No significant correlation was found between S and mass-specific RMR among species. As predicted, a significantly negative relationship exists between S and b R among species. The results suggest that the varying b R could be attributed to cell size differences among species, as those with larger cells may face stronger surface boundary limits, as predicted by the MLB hypothesis. This mechanism represents an additional way of relating the MLB and the CM hypotheses and does not exclude another mechanism based on the recent contextual multimodal theory.

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