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The response of mature forest ecosystems to a rising atmospheric carbon dioxide concentration (Ca) is a major uncertainty in projecting the future trajectory of the Earth's climate. Although leaf-level net photosynthesis is typically stimulated by exposure to elevated Ca (eCa), it is unclear how this stimulation translates into carbon cycle responses at the ecosystem scale. Here we estimate a key component of the carbon cycle, the gross primary productivity (GPP), of a mature native eucalypt forest exposed to free-air CO2 enrichment (the EucFACE experiment). In this experiment, light-saturated leaf photosynthesis increased by 19 % in response to a 38 % increase in Ca. We used the process-based forest canopy model, MAESPA, to upscale these leaf-level measurements of photosynthesis with canopy structure to estimate the GPP and its response to eCa. We assessed the direct impact of eCa, as well as the indirect effect of photosynthetic acclimation to eCa and variability among treatment plots using different model scenarios. At the canopy scale, MAESPA estimated a GPP of 1574 g C m−2 yr−1 under ambient conditions across 4 years and a direct increase in the GPP of +11 % in response to eCa. The smaller canopy-scale response simulated by the model, as compared with the leaf-level response, could be attributed to the prevalence of RuBP regeneration limitation of leaf photosynthesis within the canopy. Photosynthetic acclimation reduced this estimated response to 10 %. After taking the baseline variability in the leaf area index across plots in account, we estimated a field GPP response to eCa of 6 % with a 95 % confidence interval (−2 %, 14 %). These findings highlight that the GPP response of mature forests to eCa is likely to be considerably lower than the response of light-saturated leaf photosynthesis. Our results provide an important context for interpreting the eCa responses of other components of the ecosystem carbon cycle.

Rising atmospheric CO2 concentrations is expected to stimulate photosynthesis and carbohydrate production, while inhibiting photorespiration. By contrast, nitrogen (N) concentrations in leaves generally tend to decline under elevated CO2 (eCO2 ), which may reduce the magnitude of photosynthetic enhancement. We tested two hypotheses as to why leaf N is reduced under eCO2 : (a) A "dilution effect" caused by increased concentration of leaf carbohydrates; and (b) inhibited nitrate assimilation caused by reduced supply of reductant from photorespiration under eCO2 . This second hypothesis is fully tested in the field for the first time here, using tall trees of a mature Eucalyptus forest exposed to Free-Air CO2 Enrichment (EucFACE) for five years. Fully expanded young and mature leaves were both measured for net photosynthesis, photorespiration, total leaf N, nitrate ( NO3- ) concentrations, carbohydrates and NO3- reductase activity to test these hypotheses. Foliar N concentrations declined by 8% under eCO2 in new leaves, while the NO3- fraction and total carbohydrate concentrations remained unchanged by CO2 treatment for either new or mature leaves. Photorespiration decreased 31% under eCO2 supplying less reductant, and in situ NO3- reductase activity was concurrently reduced (-34%) in eCO2 , especially in new leaves during summer periods. Hence, NO3- assimilation was inhibited in leaves of E. tereticornis and the evidence did not support a significant dilution effect as a contributor to the observed reductions in leaf N concentration. This finding suggests that the reduction of NO3- reductase activity due to lower photorespiration in eCO2 can contribute to understanding how eCO2 -induced photosynthetic enhancement may be lower than previously expected. We suggest that large-scale vegetation models simulating effects of eCO2 on N biogeochemistry include both mechanisms, especially where NO3- is major N source to the dominant vegetation and where leaf flushing and emergence occur in temperatures that promote high photorespiration rates.

Atmospheric carbon dioxide enrichment (eCO2) can enhance plant carbon uptake and growth1–5, thereby providing an important negative feedback to climate change by slowing the rate of increase of the atmospheric CO2 concentration6. Although evidence gathered from young aggrading forests has generally indicated a strong CO2 fertilization effect on biomass growth3–5, it is unclear whether mature forests respond to eCO2 in a similar way. In mature trees and forest stands7–10, photosynthetic uptake has been found to increase under eCO2 without any apparent accompanying growth response, leaving the fate of additional carbon fixed under eCO2 unclear4,5,7–11. Here using data from the first ecosystem-scale Free-Air CO2 Enrichment (FACE) experiment in a mature forest, we constructed a comprehensive ecosystem carbon budget to track the fate of carbon as the forest responded to four years of eCO2 exposure. We show that, although the eCO2 treatment of +150 parts per million (+38 per cent) above ambient levels induced a 12 per cent (+247 grams of carbon per square metre per year) increase in carbon uptake through gross primary production, this additional carbon uptake did not lead to increased carbon sequestration at the ecosystem level. Instead, the majority of the extra carbon was emitted back into the atmosphere via several respiratory fluxes, with increased soil respiration alone accounting for half of the total uptake surplus. Our results call into question the predominant thinking that the capacity of forests to act as carbon sinks will be generally enhanced under eCO2, and challenge the efficacy of climate mitigation strategies that rely on ubiquitous CO2 fertilization as a driver of increased carbon sinks in global forests. Carbon dioxide enrichment of a mature forest resulted in the emission of the excess carbon back into the atmosphere via enhanced ecosystem respiration, suggesting that mature forests may be limited in their capacity to mitigate climate change.

Elevated CO2 affects C cycling processes which in turn can influence the nitrogen (N) and phosphorus (P) concentrations of plant tissues. Given differences in how N and P are used by plants, we asked if their stoichiometry in leaves and wood was maintained or altered in a long-term elevated CO2 experiment in a mature Eucalyptus forest on a low P soil (EucFACE). We measured N and P concentrations in green leaves at different ages at the top of mature trees across six years including five years in elevated CO2. N and P concentrations in green and senesced leaves and wood were determined to evaluate both spatial and temporal variation of leaf N and P concentrations, including the N and P retranslocation in leaves and wood. Leaf P concentrations were 32% lower in old mature leaves compared to newly flushed leaves with no effect of elevated CO2 on leaf P. By contrast, elevated CO2 significantly decreased leaf N concentrations in newly-flushed leaves but this effect disappeared as leaves matured. As such, newly-flushed leaves had 9% lower N:P ratios in elevated CO2 and N:P ratios were not different in mature green leaves (CO2 by Age effect, P = 0.02). Over time, leaf N and P concentrations in the upper canopy slightly declined in both CO2 treatments compared to before the start of the experiment. P retranslocation in leaves was 50%, almost double that of N retranslocation (29%), indicating that this site was P-limited and that P retranslocation was an important mechanism in this ecosystem to retain P in plants. As P-limited trees tend to store relatively more N than P, we found an increased N:P ratio in sapwood in response to elevated CO2 (P < 0.01), implying N accumulation in live wood. The flexible stoichiometric ratios we observed can have important implications for how plants adjust to variable environmental conditions including climate change. Hence, variable nutrient stoichiometry should be accounted for in large-scale Earth Systems models invoking biogeochemical processes.

Climate change can negatively affect sensitive tree species, affecting their acclimation and adaptation strategies. A common garden experiment provides an opportunity to test whether responses of trees from different provenances are genetically driven and if this response is related to factors at the site of origin. We hypothesized that antioxidative defence systems and leaf mass area ofAcacia melanoxylonR. Br. samples collected from different provenances will vary depending on local rainfall. Thirteen provenances ofA. melanoxylonoriginating from different rainfall habitats (500-2000 mm) were grown for 5 years in a common garden. For 2 years, phyllode samples were collected during winter and summer, for measurements of leaf mass area and concentrations of glutathione and ascorbic acid. Leaf mass area varied between seasons, years and provenances ofA. melanoxylon, and an increase was associated with decreasing rainfall at the site of origin. Ascorbic acid and glutathione concentrations varied between seasons, years (i.e., environmental factors) and among provenances ofA. melanoxylon In general, glutathione and ascorbic acid concentrations were higher in winter compared with summer. Ascorbic acid and glutathione were different among provenances, but this was not associated with rainfall at the site of origin.

Only a few previous studies have examined how photosynthetically active radiation absorptance, pigments and electron flow change in mature trees exposed to long-term increase in CO2 concentration. We investigated pigment concentrations, leaf optical properties and quantum yield of old and new leaves exposed to ambient (aCO2) and elevated (eCO2) CO2 treatments. Leaf absorptance was around 90% in E. tereticornis trees across both foliage age classes and CO2 treatments. New leaves had 15% higher quantum yield with increased absorptance within the blue spectrum than old leaves; while they reflected and transmitted more photons. In addition, young foliage had increased mass-based concentrations of chlorophyll and carotenoids; however, pigment concentrations were reduced when expressed on area-basis. Quantum yield was 9% higher in eCO2 than aCO2 across both foliage age classes. The CO2 effect was stronger in new leaves where the quantum yield was 17% higher in eCO2 than aCO2, but not different in old leaves between CO2 treatments. New leaves had higher transmittance of photons in eCO2 than aCO2, while there was no change in old leaves. Mass-based concentrations of chlorophyll and carotenoids were reduced in eCO2 compared to aCO2 while concentrations of anthocyanins were higher in response to CO2 treatment. There was a significant effect of Age x CO2 interaction on ratio a/b with larger eCO2-related reductions in old leaves (–5%) but no change in new leaves. Generally, new leaves were more efficient in utilizing the absorbed photons than old leaves, especially under eCO2 which resulted in more carbon fixation. This implies that leaves can adjust their light harvesting capacity to eCO2, particularly in younger leaves which have higher photosynthetic activity.