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Whether the terrestrial biosphere will continue to act as a net carbon (C) sink in the face of multiple global changes is questionable. A key uncertainty is whether increases in plant C fixation under elevated carbon dioxide (CO2) will translate into decades-long C storage and whether this depends on other concurrently changing factors. We investigated how manipulations of CO2, soil nitrogen (N) supply, and plant species richness influenced total ecosystem (plant + soil to 60 cm) C storage over 19 y in a free-air CO2 enrichment grassland experiment (BioCON) in Minnesota. On average, after 19 y of treatments, increasing species richness from 1 to 4, 9, or 16 enhanced total ecosystem C storage by 22 to 32%, whereas N addition of 4 g N m-2 · y-1 and elevated CO2 of +180 ppm had only modest effects (increasing C stores by less than 5%). While all treatments increased net primary productivity, only increasing species richness enhanced net primary productivity sufficiently to more than offset enhanced C losses and substantially increase ecosystem C pools. Effects of the three global change treatments were generally additive, and we did not observe any interactions between CO2 and N. Overall, our results call into question whether elevated CO2 will increase the soil C sink in grassland ecosystems, helping to slow climate change, and suggest that losses of biodiversity may influence C storage as much as or more than increasing CO2 or high rates of N deposition in perennial grassland systems. [ABSTRACT FROM AUTHOR]

Global changes can interact to affect photosynthesis and thus ecosystem carbon capture, yet few multi‐factor field studies exist to examine such interactions. Here, we evaluate leaf gas exchange responses of five perennial grassland species from four functional groups to individual and interactive global changes in an open‐air experiment in Minnesota, USA, including elevated CO2 (eCO2), warming, reduced rainfall and increased soil nitrogen supply. All four factors influenced leaf net photosynthesis and/or stomatal conductance, but almost all effects were context‐dependent, i.e. they differed among species, varied with levels of other treatments and/or depended on environmental conditions. Firstly, the response of photosynthesis to eCO2 depended on species and nitrogen, became more positive as vapour pressure deficit increased and, for a C4 grass and a legume, was more positive under reduced rainfall. Secondly, reduced rainfall increased photosynthesis in three functionally distinct species, potentially via acclimation to low soil moisture. Thirdly, warming had positive, neutral or negative effects on photosynthesis depending on species and rainfall. Overall, our results show that interactions among global changes and environmental conditions may complicate predictions based on simple theoretical expectations of main effects, and that the factors and interactions influencing photosynthesis vary among herbaceous species. Effects of global changes – elevated CO2, warming, reduced rainfall and increased soil nitrogen supply – on leaf gas exchange were largely context‐dependent, as they differed among species, varied across levels of other treatments and/or depended on environmental conditions. [ABSTRACT FROM AUTHOR]

Uncertainty about long‐term leaf‐level responses to atmospheric CO2 rise 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 eCO2 on rates of net photosynthesis and stomatal conductance in 14 perennial grassland species from four functional groups over two decades in a Minnesota Free‐Air CO2 Enrichment experiment, BioCON. Monocultures of species belonging to C3 grasses, C4 grasses, forbs, and legumes were exposed to two levels of CO2 and nitrogen supply in factorial combinations over 21 years. eCO2 increased photosynthesis by 12.9% on average in C3 species, 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 eCO2 was observed beginning in the first year and did not strengthen through time. Yet, contrary to expectations, the response of photosynthesis to eCO2 was not enhanced by increased nitrogen supply. Differences in responses among herbaceous plant functional groups were modest, with legumes responding the most and C4 grasses the least as expected, but did not further diverge over time. Leaf‐level water‐use efficiency increased by 50% under eCO2 primarily because of reduced stomatal conductance. Our results imply that enhanced nitrogen supply will not necessarily diminish photosynthetic acclimation to eCO2 in nitrogen‐limited systems, and that significant and consistent declines in stomatal conductance and increases in water‐use efficiency under eCO2 may allow plants to better withstand drought. [ABSTRACT FROM AUTHOR]

Biogeochemical models that incorporate nitrogen (N) limitation indicate that N availability will control the magnitude of ecosystem carbon uptake in response to rising CO2. Some models, however, suggest that elevated CO2 may promote ecosystem N accumulation, a feedback that in the long term could circumvent N limitation of the CO2 response while mitigating N pollution. We tested this prediction using a nine-year CO2xN experiment in a tidal marsh. Although the effects of CO2 are similar between uplands and wetlands in many respects, this experiment offers a greater likelihood of detecting CO2 effects on N retention on a decadal timescale because tidal marshes have a relatively open N cycle and can accrue soil organic matter rapidly. To determine how elevated CO2 affects N dynamics, we assessed the three primary fates of N in a tidal marsh: (1) retention in plants and soil, (2) denitrification to the atmosphere, and (3) tidal export. We assessed changes in N pools and tracked the fate of a 15N tracer added to each plot in 2006 to quantify the fraction of added N retained in vegetation and soil, and to estimate lateral N movement. Elevated CO2 alone did not increase plant N mass, soil N mass, or 15N label retention. Unexpectedly, CO2 and N interacted such that the combined N+CO2 treatment increased ecosystem N accumulation despite the stimulation in N losses indicated by reduced 15N label retention. These findings suggest that in N-limited ecosystems, elevated CO2 is unlikely to increase long-term N accumulation and circumvent progressive N limitation without additional N inputs, which may relieve plant-microbe competition and allow for increased plant N uptake. [ABSTRACT FROM AUTHOR]

Global changes can interact to affect photosynthesis and thus ecosystem carbon capture, yet few multi-factor field studies exist to examine such interactions. Here, we evaluate leaf gas exchange responses of five perennial grassland species from four functional groups to individual and interactive global changes in an open-air experiment in Minnesota, USA, including elevated CO 2 (eCO 2 ), warming, reduced rainfall and increased soil nitrogen supply. All four factors influenced leaf net photosynthesis and/or stomatal conductance, but almost all effects were context-dependent, i.e. they differed among species, varied with levels of other treatments and/or depended on environmental conditions. Firstly, the response of photosynthesis to eCO 2 depended on species and nitrogen, became more positive as vapour pressure deficit increased and, for a C 4 grass and a legume, was more positive under reduced rainfall. Secondly, reduced rainfall increased photosynthesis in three functionally distinct species, potentially via acclimation to low soil moisture. Thirdly, warming had positive, neutral or negative effects on photosynthesis depending on species and rainfall. Overall, our results show that interactions among global changes and environmental conditions may complicate predictions based on simple theoretical expectations of main effects, and that the factors and interactions influencing photosynthesis vary among herbaceous species., (© 2020 John Wiley & Sons Ltd.)

Uncertainty about long-term leaf-level responses to atmospheric CO 2 rise is a major knowledge gap that exists because of limited empirical data. Thus, it remains unclear how responses of leaf gas exchange to elevated CO 2 (eCO 2 ) 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 eCO 2 on rates of net photosynthesis and stomatal conductance in 14 perennial grassland species from four functional groups over two decades in a Minnesota Free-Air CO 2 Enrichment experiment, BioCON. Monocultures of species belonging to C 3 grasses, C 4 grasses, forbs, and legumes were exposed to two levels of CO 2 and nitrogen supply in factorial combinations over 21 years. eCO 2 increased photosynthesis by 12.9% on average in C 3 species, 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 eCO 2 was observed beginning in the first year and did not strengthen through time. Yet, contrary to expectations, the response of photosynthesis to eCO 2 was not enhanced by increased nitrogen supply. Differences in responses among herbaceous plant functional groups were modest, with legumes responding the most and C 4 grasses the least as expected, but did not further diverge over time. Leaf-level water-use efficiency increased by 50% under eCO 2 primarily because of reduced stomatal conductance. Our results imply that enhanced nitrogen supply will not necessarily diminish photosynthetic acclimation to eCO 2 in nitrogen-limited systems, and that significant and consistent declines in stomatal conductance and increases in water-use efficiency under eCO 2 may allow plants to better withstand drought., (© 2019 John Wiley & Sons Ltd.)

Biogeochemical models that incorporate nitrogen (N) limitation indicate that N availability will control the magnitude of ecosystem carbon uptake in response to rising CO2 . Some models, however, suggest that elevated CO2 may promote ecosystem N accumulation, a feedback that in the long term could circumvent N limitation of the CO2 response while mitigating N pollution. We tested this prediction using a nine-year CO2 xN experiment in a tidal marsh. Although the effects of CO2 are similar between uplands and wetlands in many respects, this experiment offers a greater likelihood of detecting CO2 effects on N retention on a decadal timescale because tidal marshes have a relatively open N cycle and can accrue soil organic matter rapidly. To determine how elevated CO2 affects N dynamics, we assessed the three primary fates of N in a tidal marsh: (1) retention in plants and soil, (2) denitrification to the atmosphere, and (3) tidal export. We assessed changes in N pools and tracked the fate of a (15) N tracer added to each plot in 2006 to quantify the fraction of added N retained in vegetation and soil, and to estimate lateral N movement. Elevated CO2 alone did not increase plant N mass, soil N mass, or (15) N label retention. Unexpectedly, CO2 and N interacted such that the combined N+CO2 treatment increased ecosystem N accumulation despite the stimulation in N losses indicated by reduced (15) N label retention. These findings suggest that in N-limited ecosystems, elevated CO2 is unlikely to increase long-term N accumulation and circumvent progressive N limitation without additional N inputs, which may relieve plant-microbe competition and allow for increased plant N uptake., (© 2015 John Wiley & Sons Ltd.)