Does Bertrand's rule apply to macronutrients?

It has been known for over a century that the dose-response curve for many micronutrients is non-monotonic, having an initial stage of increasing benefits with increased intake, followed by increasing costs as excesses become toxic. This phenomenon, termed Bertrand's rule, is widely assumed not to apply to caloric macronutrients. To date this assumption has been safe, owing to the considerable methodological challenges involved in coaxing animals to over-ingest macronutrients in a way that enables the effects of specific food components to be isolated. Here we report an experiment which overcomes these difficulties, to test whether the second phase (incurring costs with excessive intake) applies to carbohydrate intake by the generalist-feeding caterpillar Spodoptera littoralis. The results showed that excess carbohydrate intake caused increased mortality, thus extending Bertrand's rule to macronutrients.