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Summary Natural selection favors growth by selecting a combination of plant traits that maximize photosynthetic CO₂assimilation at the lowest combined carbon costs of resource acquisition and use. We quantified how soil nutrient availability, plant nutrient acquisition strategies, and aridity modulate the variability in plant costs of nutrient acquisition relative to water acquisition (β).We used an eco‐evolutionary optimality framework and a global carbon isotope dataset to quantify β.Under low soil nitrogen‐to‐carbon (N : C) ratios, a mining strategy (symbioses with ectomycorrhizal and ericoid mycorrhizal fungi) reduced β by mining organic nitrogen, compared with a scavenging strategy (symbioses with arbuscular mycorrhizal fungi). Conversely, under high N : C ratios, scavenging strategies reduced β by effectively scavenging soluble nitrogen, compared with mining strategies. N₂‐fixing plants did not exhibit reduced β under low N : C ratios compared with non‐N₂‐fixing plants. Moisture increased β only in plants using a scavenging strategy, reflecting direct impacts of aridity on the carbon costs of maintaining transpiration in these plants. Nitrogen and phosphorus colimitation further modulated β.Our findings provide a framework for simulating the variability of plant economics due to plant nutrient acquisition strategies in earth system models. 

ABSTRACT 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 amongst 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 the largest in environments where demand is the greatest. Using a nutrient addition experiment replicated across 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 the highest theoretical leaf nitrogen demand, increasing more in colder and drier environments than warmer and wetter environments. Thus, leaf nitrogen responses to nitrogen supply are primarily influenced by climatic gradients in photosynthetic nitrogen demand, an insight that could improve ESM predictions. 

Summary Interactions between carbon (C) and nitrogen (N) cycles in terrestrial ecosystems are simulated in advanced vegetation models, yet methodologies vary widely, leading to divergent simulations of past land C balance trends. This underscores the need to reassess our understanding of ecosystem processes, given recent theoretical advancements and empirical data. We review current knowledge, emphasising evidence from experiments and trait data compilations for vegetation responses to CO₂and N input, alongside theoretical and ecological principles for modelling. N fertilisation increases leaf N content but inconsistently enhances leaf‐level photosynthetic capacity. Whole‐plant responses include increased leaf area and biomass, with reduced root allocation and increased aboveground biomass. Elevated atmospheric CO₂also boosts leaf area and biomass but intensifies belowground allocation, depleting soil N and likely reducing N losses. Global leaf traits data confirm these findings, indicating that soil N availability influences leaf N content more than photosynthetic capacity. A demonstration model based on the functional balance hypothesis accurately predicts responses to N and CO₂fertilisation on tissue allocation, growth and biomass, offering a path to reduce uncertainty in global C cycle projections.