Resource limitation in a competitive context determines complex plant responses to experimental resource additions

Almost all models of plant resource limitation are grounded in either one or both of two simple conceptual models: Liebig's Minimum Hypothesis (LMH), the idea that plants are limited by the resource in shortest supply, and the Multiple Limitation Hypothesis (MLH), the idea that plants should adjust to their environment so that all essential resources are equally limiting. Despite the differences in their predictions, experiments have so far failed to discriminate between them. In a simple factorial nitrogen and water addition experiment in a Minnesota grassland, we observed shifts in allocation that, as in previous studies, are not all explained by a single theory. We found that leaf biomass responded positively to nitrogen additions but did not respond to water additions. We found that fine-root biomass increased in response to water additions, but only at low nitrogen levels, and that fine-root biomass decreased in response to nitrogen additions, but only at high water levels. To understand these responses we built a physiologically based model of plant competition for water, nitrogen, and space to predict plant allocation to fine roots and leaves. Critically, we include in our model the inherent variability of soil moisture and treat light, water, and nitrogen as resources with distinct mechanistic roles. Experimental results showed that plants were nitrogen and water limited. The model explains the experimental results, under conditions of co-limitation, as follows. Foliage increases with nitrogen additions but not water additions because leaf construction is constrained by nitrogen uptake. When water is added, plants spend a larger fraction of the growing season limited by light (and effectively nitrogen) than by water. Thus, water additions cause fine-root biomass to increase because of the increased importance of nitrogen limitation. The response of fine-root biomass to water additions decreases with nitrogen additions because these additions reduce nitrogen limitation. In general, our results are explained by sequential resource limitation. The rate of carbon assimilation may be limited by a single resource at any one moment, but the identity of the limiting resource(s) changes throughout the growing season.