Accurately representing the relationships between nitrogen supply and photosynthesis is crucial for reliably predicting carbon-nitrogen cycle coupling in Earth System Models (ESMs). Most ESMs assume positive correlations among soil nitrogen supply, leaf nitrogen content, and photosynthetic capacity. However, leaf photosynthetic nitrogen demand may influence the leaf nitrogen response to soil nitrogen supply, thus responses to nitrogen supply are expected to be largest in environments where demand is greatest. Using a nutrient addition experiment replicated at 26 sites spanning four continents, we demonstrated that climate variables were stronger predictors of leaf nitrogen content than soil nutrient supply. Leaf nitrogen increased more strongly with soil nitrogen supply in regions with highest theoretical leaf nitrogen demand, increasing more in colder and drier environments than warmer and wetter ones. Thus, leaf nitrogen responses to nitrogen supply are primarily influenced by climatic gradients in photosynthetic nitrogen demand, an insight that will improve ESM predictions.

Empirical studies have shown that plant photosynthetic responses to environmental change can vary over time due to acclimation, but acclimation responses are often not included in Earth System Models (ESMs). Photosynthetic least cost theory can be used to develop models of photosynthetic acclimation that are simple and testable. The theory is based on the idea that, optimally, plants will acclimate to maintain the fastest rate of photosynthesis at the lowest water and nutrient use. Formulations of this theory have been developed for C3 plants, but not C4 plants, which account for over 20% global photosynthesis and are over-represented among widely grown crops. Here, we use photosynthetic least cost theory to derive a model for C4 photosynthetic acclimation to above-ground abiotic conditions. We then compare our model’s responses to a similar model of C3 photosynthetic acclimation and find that C4 photosynthesis has the highest simulated advantage over C3 photosynthesis in hot and low CO2 environments. We find that this advantage predicts C4 abundance globally, but that the shallower CO2 response of C4 as compared to C3 photosynthesis will reduce C4 plant competitiveness under future conditions, despite higher temperatures. We also show that an acclimated model predicts similar or faster rates of C4 under all conditions than a model that does not consider acclimation, suggesting that ESMs are underestimating future C4 carbon uptake by not including acclimation. Our model is designed for easy incorporation into such ESMs.

Despite widespread evidence that biological invasion influences both the biotic and abiotic soil environments, the extent to which these two pathways underpin the effects of invasion on plant traits and performance is unknown. Leveraging a long-term (14-yr) field experiment, we show that an allelochemical-producing invader affects plants through biotic mechanisms, altering the soil fungal community composition, with no apparent shifts in soil nutrient availability. Changes in belowground fungal communities result in high costs of nutrient uptake for native perennials and a shift in functional traits linked to their water and nutrient use efficiencies. Some species in the invaded community compensate for high nutrient costs by reducing nutrient uptake and maintaining photosynthesis by expending more water, which demonstrates a trade-off in trait investment. For the first time, we show that the disruption of belowground nutritional symbionts can drive native plants toward novel regions in order to maintain their water and nutrient economics.