When Michaelis and Menten met Holling: towards a mechanistic theory of plant nutrient foraging behaviour.

Plants are adept at assessing and responding to nutrients in soil, and generally proliferate roots into nutrient-rich patches. An analogy between this growth response and animal foraging movement is often drawn, but because of differences between plants and animals it has not always been clear how to directly apply existing foraging theory to plants. Here we suggest one way to unite pre-existing ideas in plant nutrient uptake with foraging theory. First, we show that the Michaelis-Menten equation used by botanists and the Holling disc equation used by zoologists are actually just rearrangements of the same functional response. This mathematical unity permits the translation of existing knowledge about the nutrient uptake physiology of plants into the language of foraging behaviour, and as a result gives botanists direct access to foraging theory. Second, we developed a model of root foraging precision based on the Holling disc equation and the marginal value theorem, and parameterize it from the literature. The model predicts (i) generally plants should invest in higher quality patches compared to lower quality patches, and as patch background-contrast increases; (ii) low encounter rates between roots and nutrients result in high root foraging precision; and (iii) low handling times for nutrients should result in high root foraging precision. The available data qualitatively support these predictions. Third, to parameterize the model above we undertook a review of the literature. From that review we obtained parameter estimates for nitrate and/or ammonium uptake for 45 plant species from 38 studies. We observe that the parameters ranged over six orders of magnitude, there was no trade-off in foraging ability for nitrate versus ammonium: plants that were efficient foragers for one form of nitrogen were efficient foragers for the other, and there was also no phylogenetic signal in the parameter estimates.