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Abstract Plants respond to increasing atmospheric CO2 concentrations by reducing leaf nitrogen content and photosynthetic capacity—patterns that correspond with increased net photosynthesis and growth. Despite the longstanding notion that nitrogen availability regulates these responses, eco-evolutionary optimality theory posits that leaf-level responses to elevated CO2 are driven by leaf nitrogen demand for building and maintaining photosynthetic enzymes and are independent of nitrogen availability. In this study, we examined leaf and whole-plant responses of Glycine max L. (Merr) subjected to full-factorial combinations of two CO2, two inoculation, and nine nitrogen fertilization treatments. Nitrogen fertilization and inoculation did not alter leaf photosynthetic responses to elevated CO2. Instead, elevated CO2 decreased the maximum rate of ribulose-1,5-bisophosphate oxygenase/carboxylase (Rubisco) carboxylation more strongly than it decreased the maximum rate of electron transport for ribulose-1,5-bisphosphate (RuBP) regeneration, increasing net photosynthesis by allowing rate-limiting steps to approach optimal coordination. Increasing fertilization enhanced positive whole-plant responses to elevated CO2 due to increased below-ground carbon allocation and nitrogen uptake. Inoculation with nitrogen-fixing bacteria did not influence plant responses to elevated CO2. These results reconcile the role of nitrogen availability in plant responses to elevated CO2, showing that leaf photosynthetic responses are regulated by leaf nitrogen demand while whole-plant responses are constrained by nitrogen availability. 

Summary Affecting biodiversity, plants with larger genome sizes (GS) may be restricted in nutrient‐poor conditions. This pattern has been attributed to their greater cellular nitrogen (N) and phosphorus (P) investments and hypothesized nutrient–investment tradeoffs between cell synthesis and physiological attributes associated with growth. However, the influence of GS on cell size and functioning may also contribute to GS‐dependent growth responses to nutrients.To test whether and how GS is associated with cellular nutrient, stomata, and/or physiological attributes, we examined > 500 forbs and grasses from seven grassland sites conducting a long‐term N and P fertilization experiment.Larger GS plants had increased cellular nutrient contents and larger, but fewer stomata than smaller GS plants. Larger GS grasses (but not forbs) also had lower photosynthetic rates and water‐use efficiencies. However, nutrients had no direct effect on GS‐dependent physiological attributes and GS‐dependent physiological changes likely arise from how GS influences cells. At the driest sites, large GS grasses displayed high water‐use efficiency mostly because transpiration was reduced relative to photosynthesis in these conditions.We suggest that climatic conditions and GS‐associated cell traits that modify physiological responses, rather than resource–investment tradeoffs, largely explain GS‐dependent growth responses to nutrients (especially for grasses). 

Abstract The connection between soil nitrogen availability, leaf nitrogen, and photosynthetic capacity is not perfectly understood. Because these three components tend to be positively related over large spatial scales, some posit that soil nitrogen positively drives leaf nitrogen, which positively drives photosynthetic capacity. Alternatively, others posit that photosynthetic capacity is primarily driven by above-ground conditions. Here, we examined the physiological responses of a non-nitrogen-fixing plant (Gossypium hirsutum) and a nitrogen-fixing plant (Glycine max) in a fully factorial combination of light by soil nitrogen availability to help reconcile these competing hypotheses. Soil nitrogen stimulated leaf nitrogen in both species, but the relative proportion of leaf nitrogen used for photosynthetic processes was reduced under elevated soil nitrogen in all light availability treatments due to greater increases in leaf nitrogen content than chlorophyll and leaf biochemical process rates. Leaf nitrogen content and biochemical process rates in G. hirsutum were more responsive to changes in soil nitrogen than those in G. max, probably due to strong G. max investments in root nodulation under low soil nitrogen. Nonetheless, whole-plant growth was significantly enhanced by increased soil nitrogen in both species. Light availability consistently increased relative leaf nitrogen allocation to leaf photosynthesis and whole-plant growth, a pattern that was similar between species. These results suggest that the leaf nitrogen–photosynthesis relationship varies under different soil nitrogen levels and that these species preferentially allocated more nitrogen to plant growth and non-photosynthetic leaf processes, rather than photosynthesis, as soil nitrogen increased. 

Abstract Forbs (“wildflowers”) are important contributors to grassland biodiversity but are vulnerable to environmental changes. In a factorial experiment at 94 sites on 6 continents, we test the global generality of several broad predictions: (1) Forb cover and richness decline under nutrient enrichment, particularly nitrogen enrichment. (2) Forb cover and richness increase under herbivory by large mammals. (3) Forb richness and cover are less affected by nutrient enrichment and herbivory in more arid climates, because water limitation reduces the impacts of competition with grasses. (4) Forb families will respond differently to nutrient enrichment and mammalian herbivory due to differences in nutrient requirements. We find strong evidence for the first, partial support for the second, no support for the third, and support for the fourth prediction. Our results underscore that anthropogenic nitrogen addition is a major threat to grassland forbs, but grazing under high herbivore intensity can offset these nutrient effects. 

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. 

Summary Allocation of leaf phosphorus (P) among different functional fractions represents a crucial adaptive strategy for optimizing P use. However, it remains challenging to monitor the variability in leaf P fractions and, ultimately, to understand P‐use strategies across diverse plant communities.We explored relationships between five leaf P fractions (orthophosphate P, P_i; lipid P, P_L; nucleic acid P, P_N; metabolite P, P_M; and residual P, P_R) and 11 leaf economic traits of 58 woody species from three biomes in China, including temperate, subtropical and tropical forests. Then, we developed trait‐based models and spectral models for leaf P fractions and compared their predictive abilities.We found that plants exhibiting conservative strategies increased the proportions of P_Nand P_M, but decreased the proportions of P_iand P_L, thus enhancing photosynthetic P‐use efficiency, especially under P limitation. Spectral models outperformed trait‐based models in predicting cross‐site leaf P fractions, regardless of concentrations (R² = 0.50–0.88 vs 0.34–0.74) or proportions (R² = 0.43–0.70 vs 0.06–0.45).These findings enhance our understanding of leaf P‐allocation strategies and highlight reflectance spectroscopy as a promising alternative for characterizing large‐scale leaf P fractions and plant P‐use strategies, which could ultimately improve the physiological representation of the plant P cycle in land surface models. 

Summary Leaf dark respiration (R_d) acclimates to environmental changes. However, the magnitude, controls and time scales of acclimation remain unclear and are inconsistently treated in ecosystem models.We hypothesized thatR_dand Rubisco carboxylation capacity (V_cmax) at 25°C (R_d,25,V_cmax,25) are coordinated so thatR_d,25variations supportV_cmax,25at a level allowing full light use, withV_cmax,25reflecting daytime conditions (for photosynthesis), andR_d,25/V_cmax,25reflecting night‐time conditions (for starch degradation and sucrose export). We tested this hypothesis temporally using a 5‐yr warming experiment, and spatially using an extensive field‐measurement data set. We compared the results to three published alternatives:R_d,25declines linearly with daily average prior temperature;R_dat average prior night temperatures tends towards a constant value; andR_d,25/V_cmax,25is constant.Our hypothesis accounted for more variation in observedR_d,25over time (R² = 0.74) and space (R² = 0.68) than the alternatives. Night‐time temperature dominated the seasonal time‐course ofR_d, with an apparent response time scale ofc.2 wk.V_cmaxdominated the spatial patterns.Our acclimation hypothesis results in a smaller increase in globalR_din response to rising CO₂and warming than is projected by the two of three alternative hypotheses, and by current models. 

Abstract Dominance often indicates one or a few species being best suited for resource capture and retention in a given environment. Press perturbations that change availability of limiting resources can restructure competitive hierarchies, allowing new species to capture or retain resources and leaving once dominant species fated to decline. However, dominant species may maintain high abundances even when their new environments no longer favour them due to stochastic processes associated with their high abundance, impeding deterministic processes that would otherwise diminish them.Here, we quantify the persistence of dominance by tracking the rate of decline in dominant species at 90 globally distributed grassland sites under experimentally elevated soil nutrient supply and reduced vertebrate consumer pressure.We found that chronic experimental nutrient addition and vertebrate exclusion caused certain subsets of species to lose dominance more quickly than in control plots. In control plots, perennial species and species with high initial cover maintained dominance for longer than annual species and those with low initial cover respectively. In fertilized plots, species with high initial cover maintained dominance at similar rates to control plots, while those with lower initial cover lost dominance even faster than similar species in controls. High initial cover increased the estimated time to dominance loss more strongly in plots with vertebrate exclosures than in controls. Vertebrate exclosures caused a slight decrease in the persistence of dominance for perennials, while fertilization brought perennials' rate of dominance loss in line with those of annuals. Annual species lost dominance at similar rates regardless of treatments.Synthesis.Collectively, these results point to a strong role of a species' historical abundance in maintaining dominance following environmental perturbations. Because dominant species play an outsized role in driving ecosystem processes, their ability to remain dominant—regardless of environmental conditions—is critical to anticipating expected rates of change in the structure and function of grasslands. Species that maintain dominance while no longer competitively favoured following press perturbations due to their historical abundances may result in community compositions that do not maximize resource capture, a key process of system responses to global change.