Too much of a good thing: on stoichiometrically balanced diets and maximal growth

Nutritional imbalances are of great interest in the ecological stoichiometry literature, in which researchers have focused almost exclusively on cases where nutrients are available in low amounts relative to energy (carbon), and animal growth is impaired due to insufficient nutrient intake. Little attention has been given to situations where food elemental content is higher than the level that satisfies animal requirements. However, most animals are strongly homeostatic with respect to the elemental composition of their body; hence they must excrete the excess of elements that are not in short supply. To date, stoichiometric theory has assumed that excretion of superfluous elements does not come with a cost and, thus, that consumption of food with surplus nutrients does not impair performance. Here we challenge this assumption, based on a compilation of several examples involving food phosphorus content that show that the performance of a wide variety of animals decreases when supplied with food containing high concentrations of (potentially) limiting nutrients. We discuss possible mechanisms for this phenomenon, and suggest that animals most vulnerable to effects of high food nutrient content are those that normally feed on low- quality (low-nutrient: C) food, and have a relatively low body nutrient content themselves, such as herbivores and detritivores.