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. 
                        more » 
                        « less   
                    This content will become publicly available on November 1, 2025
                            
                            Acclimation of Photosynthesis to CO 2 Increases Ecosystem Carbon Storage due to Leaf Nitrogen Savings
                        
                    
    
            ABSTRACT Photosynthesis is the largest flux of carbon between the atmosphere and Earth's surface and is driven by enzymes that require nitrogen, namely, ribulose‐1,5‐bisphosphate (RuBisCO). Thus, photosynthesis is a key link between the terrestrial carbon and nitrogen cycle, and the representation of this link is critical for coupled carbon‐nitrogen land surface models. Models and observations suggest that soil nitrogen availability can limit plant productivity increases under elevated CO2. Plants acclimate to elevated CO2by downregulating RuBisCO and thus nitrogen in leaves, but this acclimation response is not currently included in land surface models. Acclimation of photosynthesis to CO2can be simulated by the photosynthetic optimality theory in a way that matches observations. Here, we incorporated this theory into the land surface component of the Energy Exascale Earth System Model (ELM). We simulated land surface carbon and nitrogen processes under future elevated CO2conditions to 2100 using the RCP8.5 high emission scenario. Our simulations showed that when photosynthetic acclimation is considered, photosynthesis increases under future conditions, but maximum RuBisCO carboxylation and thus photosynthetic nitrogen demand decline. We analyzed two simulations that differed as to whether the saved nitrogen could be used in other parts of the plant. The allocation of saved leaf nitrogen to other parts of the plant led to (1) a direct alleviation of plant nitrogen limitation through reduced leaf nitrogen requirements and (2) an indirect reduction in plant nitrogen limitation through an enhancement of root growth that led to increased plant nitrogen uptake. As a result, reallocation of saved leaf nitrogen increased ecosystem carbon stocks by 50.3% in 2100 as compared to a simulation without reallocation of saved leaf nitrogen. These results suggest that land surface models may overestimate future ecosystem nitrogen limitation if they do not incorporate leaf nitrogen savings resulting from photosynthetic acclimation to elevated CO2. 
        more » 
        « less   
        
    
                            - Award ID(s):
- 2045968
- PAR ID:
- 10614024
- Publisher / Repository:
- John Wiley and Sons
- Date Published:
- Journal Name:
- Global Change Biology
- Volume:
- 30
- Issue:
- 11
- ISSN:
- 1354-1013
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
More Like this
- 
            
- 
            Abstract Uncertainty about long‐term leaf‐level responses to atmospheric CO2rise is a major knowledge gap that exists because of limited empirical data. Thus, it remains unclear how responses of leaf gas exchange to elevated CO2(eCO2) vary among plant species and functional groups, or across different levels of nutrient supply, and whether they persist over time for long‐lived perennials. Here, we report the effects of eCO2on rates of net photosynthesis and stomatal conductance in 14 perennial grassland species from four functional groups over two decades in a Minnesota Free‐Air CO2Enrichment experiment, BioCON. Monocultures of species belonging to C3grasses, C4grasses, forbs, and legumes were exposed to two levels of CO2and nitrogen supply in factorial combinations over 21 years. eCO2increased photosynthesis by 12.9% on average in C3species, substantially less than model predictions of instantaneous responses based on physiological theory and results of other studies, even those spanning multiple years. Acclimation of photosynthesis to eCO2was observed beginning in the first year and did not strengthen through time. Yet, contrary to expectations, the response of photosynthesis to eCO2was not enhanced by increased nitrogen supply. Differences in responses among herbaceous plant functional groups were modest, with legumes responding the most and C4grasses the least as expected, but did not further diverge over time. Leaf‐level water‐use efficiency increased by 50% under eCO2primarily because of reduced stomatal conductance. Our results imply that enhanced nitrogen supply will not necessarily diminish photosynthetic acclimation to eCO2in nitrogen‐limited systems, and that significant and consistent declines in stomatal conductance and increases in water‐use efficiency under eCO2may allow plants to better withstand drought.more » « less
- 
            Abstract Realistic simulation of leaf photosynthetic and respiratory processes is needed for accurate prediction of the global carbon cycle. These two processes systematically acclimate to long‐term environmental changes by adjusting photosynthetic and respiratory traits (e.g., the maximum photosynthetic capacity at 25°C (Vcmax,25) and the leaf respiration rate at 25°C (R25)) following increasingly well‐understood principles. While some land surface models (LSMs) now account for thermal acclimation, they do so by assigning empirical parameterizations for individual plant functional types (PFTs). Here, we have implemented an Eco‐Evolutionary Optimality (EEO)‐based scheme to represent the universal acclimation of photosynthesis and leaf respiration to multiple environmental effects, and that therefore requires no PFT‐specific parameterizations, in a standard version of the widely used LSM, Noah MP. We evaluated model performance with plant trait data from a 5‐year experiment and extensive global field measurements, and carbon flux measurements from FLUXNET2015. We show that observedR25andVcmax,25vary substantially both temporally and spatially within the same PFT (C.V.>20%). Our EEO‐based scheme captures 62% of the temporal and 70% of the spatial variations inVcmax,25(73% and 54% of the variations inR25). The standard scheme underestimates gross primary production by 10% versus 2% for the EEO‐based scheme and generates a larger spread inr(correlation coefficient) across flux sites (0.79 ± 0.16 vs. 0.84 ± 0.1, mean ± S.D.). The standard scheme greatly overestimates canopy respiration (bias: ∼200% vs. 8% for the EEO scheme), resulting in less CO2uptake by terrestrial ecosystems. Our approach thus simulates climate‐carbon coupling more realistically, with fewer parameters.more » « less
- 
            The efflux of carbon dioxide (CO2) from woody stems, a proxy for stem respiration, is a critical carbon flux from ecosystems to the atmosphere, which increases with temperature on short timescales. However, plants acclimate their respiratory response to temperature on longer timescales, potentially weakening the carbon-climate feedback. The magnitude of this acclimation is uncertain despite its importance for predicting future climate change. We develop an optimality-based theory dynamically linking stem respiration with leaf water supply to predict its thermal acclimation. We show that the theory accurately reproduces observations of spatial and seasonal change. We estimate the global value for current annual stem CO2efflux as 27.4 ± 5.9 PgC. By 2100, incorporating thermal acclimation reduces projected stem respiration without considering acclimation by 24 to 46%, thus reducing land ecosystem carbon emissions.more » « less
- 
            Empirical evidence and theoretical understanding of ecosystem carbon and nitrogen cycle interactionsSummary 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.more » « less
 An official website of the United States government
An official website of the United States government 
				
			 
					 
					
