Elevated carbon dioxide increases soil nitrogen and phosphorus availability in a phosphorus‐limited Eucalyptus woodland

Free‐air CO₂ enrichment (FACE) experiments have demonstrated increased plant productivity in response to elevated (e)CO₂, with the magnitude of responses related to soil nutrient status. Whilst understanding nutrient constraints on productivity responses to eCO₂ is crucial for predicting carbon uptake and storage, very little is known about how eCO₂ affects nutrient cycling in phosphorus (P)‐limited ecosystems. Our study investigates eCO₂ effects on soil N and P dynamics at the EucFACE experiment in Western Sydney over an 18‐month period. Three ambient and three eCO₂ (+150 ppm) FACE rings were installed in a P‐limited, mature Cumberland Plain Eucalyptus woodland. Levels of plant accessible nutrients, evaluated using ion exchange resins, were increased under eCO₂, compared to ambient, for nitrate (+93%), ammonium (+12%) and phosphate (+54%). There was a strong seasonality to responses, particularly for phosphate, resulting in a relatively greater stimulation in available P, compared to N, under eCO₂ in spring and summer. eCO₂ was also associated with faster nutrient turnover rates in the first six months of the experiment, with higher N (+175%) and P (+211%) mineralization rates compared to ambient rings, although this difference did not persist. Seasonally dependant effects of eCO₂ were seen for concentrations of dissolved organic carbon in soil solution (+31%), and there was also a reduction in bulk soil pH (‐0.18 units) observed under eCO₂. These results demonstrate that CO₂ fertilization increases nutrient availability – particularly for phosphate – in P‐limited soils, likely via increased plant belowground investment in labile carbon and associated enhancement of microbial turnover of organic matter and mobilization of chemically bound P. Early evidence suggests that there is the potential for the observed increases in P availability to support increased ecosystem C‐accumulation under future predicted CO₂ concentrations.     

Impacts of warming and elevated CO2 on a semi‐arid grassland are non‐additive,shift with precipitation,and reverse over time

It is unclear how elevated CO₂ (eCO₂) and the corresponding shifts in temperature and precipitation will interact to impact ecosystems over time. During a 7‐year experiment in a semi‐arid grassland, the response of plant biomass to eCO₂ and warming was largely regulated by interannual precipitation, while the response of plant community composition was more sensitive to experiment duration. The combined effects of eCO₂ and warming on aboveground plant biomass were less positive in ‘wet’ growing seasons, but total plant biomass was consistently stimulated by ~ 25% due to unique, supra‐additive responses of roots. Independent of precipitation, the combined effects of eCO₂ and warming on C₃ graminoids became increasingly positive and supra‐additive over time, reversing an initial shift toward C₄ grasses. Soil resources also responded dynamically and non‐additively to eCO₂ and warming, shaping the plant responses. Our results suggest grasslands are poised for drastic changes in function and highlight the need for long‐term, factorial experiments.     

Growth,nutrient dynamics,and efficiency responses to carbon dioxide and phosphorus nutrition in soybean

Plant mineral nutrients such as phosphorus may exert major control on crop responses to the rising atmospheric carbon dioxide (CO₂) concentrations. To evaluate the growth, nutrient dynamics, and efficiency responses to CO₂ and phosphorus nutrition, soybean (Glycine max (L.) Merr.) was grown in controlled environment growth chambers with sufficient (0.50 mM) and deficient (0.10 and 0.01 mM) phosphate (Pi) supply under ambient and elevated CO₂ (aCO₂, 400 and eCO₂, 800 µmol mol^?1, respectively). The CO₂ × Pi interaction was detected for leaf area, leaf and stem dry weight, and total plant biomass. The severe decrease in plant biomass in Pi-deficient plants (10–76%) was associated with reduced leaf area and photosynthesis (P_net). The degree of growth stimulation (0–55% total biomass) by eCO₂ was dependent upon the severity of Pi deficiency and was closely associated with the increased phosphorus utilization efficiency. With the exception of leaf and root biomass, Pi deficiency decreased the biomass partitioning to other plant organs with the maximum decrease observed in seed weight (8–42%) across CO₂ levels. The increased tissue nitrogen (N) concentration in Pi-deficient plants was accredited to the lower biomass and increased nutrient uptake due to the larger root to shoot ratio. The tissue P and N concentration tended to be lower at eCO₂ versus aCO₂ and did not appear to be the main cause of the lack of CO₂ response of growth and P_net under severe Pi deficiency. The leaf N/P ratio of >16 was detrimental to soybean growth. The tissue P concentration needed to attain the maximum productivity for biomass and seed yield tended to be higher at eCO₂ versus aCO₂. Therefore, the eCO₂ is likely to increase the leaf critical P concentration for maximum biomass productivity and yield in soybean.     

Short photoperiod attenuates CO2 fertilization effect on shoot biomass in Arabidopsis thaliana

The level of carbon dioxide (CO₂) in the air can affect several traits in plants. Elevated atmospheric CO₂ (eCO₂) can enhance photosynthesis and increase plant productivity, including biomass, although there are inconsistencies regarding the effects of eCO₂ on the plant growth response. The compounding effects of ambient environmental conditions such as light intensity, photoperiod, water availability, and soil nutrient composition can affect the extent to which eCO₂ enhances plant productivity. This study aimed to investigate the growth response of Arabidopsis thaliana to eCO₂ (800 ppm) under short photoperiod (8/16 h, light/dark cycle). Here, we report an attenuated fertilization effect of eCO₂ on the shoot biomass of Arabidopsis plants grown under short photoperiod. The biomass of two-, three-, and four-week-old Arabidopsis plants was increased by 10%, 15%, and 28%, respectively, under eCO₂ compared to the ambient CO₂ (aCO₂, 400 ppm) i.e. control. However, the number of rosette leaves, rosette area, and shoot biomass were similar in mature plants under both CO₂ conditions, despite 40% higher photosynthesis in eCO₂ exposed plants. The levels of chlorophylls and carotenoids were similar in the fully expanded rosette leaves regardless of the level of CO₂. In conclusion, CO₂ enrichment moderately increased Arabidopsis shoot biomass at the juvenile stage, whereas the eCO₂-induced increment in shoot biomass was not apparent in mature plants. A shorter day-length can limit the source-to-sink resource allocation in a plant in age-dependent manner, hence diminishing the eCO₂ fertilization effect on the shoot biomass in Arabidopsis plants grown under short photoperiod.     

Interacting effects of elevated CO2, nutrient availability and plant species on a generalist invertebrate herbivore

By affecting plant growth and phytochemistry elevated CO₂ may have indirect effects on the performance of herbivores. These effects show considerable variability across studies and may depend on nutrient availability, the carbon/nutrient‐balance in plant tissues and the secondary metabolism of plants. We studied the responses to elevated CO₂ and different nutrient availability of 12 herbaceous plant species differing in their investment into secondary compounds. Caterpillars of the generalist herbivore Spodoptera littoralis were reared on the leaves produced and their consumption and growth rates analysed. Elevated CO₂ resulted in a similar increase of biomass in all plant species, whereas the positive effect of fertilization varied among plant species. Specific leaf weight was influenced by elevated CO₂, but the effect depended on nutrient level and identity of plant species. Elevated CO₂ increased the C/N ratio of the leaves of most species. Caterpillars consumed more leaf material when plants were grown under elevated CO₂ and low nutrients. This indicates compensatory feeding due to lower tissue quality. However, the effects of elevated CO₂, nutrient availability and plant species identity on leaf consumption interacted. Both the effects of CO₂ and nutrient availability on the relative growth rate of the herbivore depended on the plant species. The feeding rate of S. littoralis on plant species that do not produce nitrogen‐containing secondary compounds (NCSC) was higher under low nutrient availability. In contrast, in plants producing NCSC nutrient availability had no effect on the feeding rate. This suggests that compensatory feeding in response to low nutrient contents may not be possible if plants produce NCSC. We conclude that elevated CO₂ causes species‐specific changes in the quality of plant tissues and consequently in changes in the preferences of herbivores for plant species. This could result in changes in plant community composition.     

Species-specific responses of plant communities to altered carbon and nutrient availability

In a field microcosm experiment, species‐specific responses of aboveground biomass of two California annual grassland communities to elevated CO₂ and nutrient availability were investigated. One community grows on shallow, nutrient‐poor serpentine‐derived soil whereas the other occurs on deeper, modestly fertile sandstone/greenstone‐derived substrate. In most species, CO₂ effects did not appear until late in the growing season, probably because the elevated CO₂ increased water‐use‐efficiency easing, the onset of the summer drought. Responses of aboveground biomass to elevated CO₂ differed depending on nutrient availability. Similarly, biomass responses to nutrient treatments differed depending on the CO₂ status. For the majority of the species, production increased most under elevated CO₂ with added nutrients (N,P,K, and micro nutrients). Some species were losers under conditions that increased overall community production, including Bromus hordeaceus in the serpentine community (negative biomass response under elevated CO₂) and Lotus wrangelianus in both communities (negative biomass response with added nitrogen). Treatment and competitive effects on species‐specific biomass varied in both magnitude and direction, especially in the serpentine community, significantly affecting community structure. Individual resource environments are likely to be affected by neighbouring plants, and these competitive interactions complicate predictions of species' responses to elevated CO₂.     

Plant productivity is a key driver of soil respiration response to climate change in a nutrient-limited soil.

Despite knowledge of the interaction between climate change factors significant uncertainty exists concerning the individual and interactive effects of elevated carbon dioxide (eCO₂) and elevated temperature (eT) on the soil microbiome and function. Here we examine the individual and interactive effects of eCO₂ and eT on tree growth, soil respiration (R_soil), biomass, structural and functional composition of microbial community, nitrogen (N) mineralisation and N availability in a whole tree chamber experiment. Eucalyptus globulus plants were grown from seedling to ca. 10 m tall for 15 months in a nutrient-poor sandy soil under ambient and elevated (+ 240 ppm) atmospheric CO₂ concentrations combined with ambient or elevated temperatures (+ 3 °C) in a full factorial design. Plant growth was strongly stimulated under eCO₂, but eT had little impact on any measured plant property. In contrast, R_soil was not consistently affected by eCO₂ or eT, but correlated strongly with root and leaf biomass. The response of N-mineralisation and nutrient availability to eCO₂ and eT varied across time, and available N correlated strongly with plant height. Further, the C:N ratio of the microbial biomass and leaves were both higher under eCeT treatment. However, these functional measures were not significantly linked to either structural or functional diversity of the soil microbiome. Taken together, these results suggest that in this low-nutrient soil, belowground processes are principally driven by aboveground productivity. Our work provides novel insight into mechanisms underlying above- and belowground response to climate change, and the potential to sequester C in a low-nutrient status soil under future climatic conditions may be limited .     

Constraints to nitrogen acquisition of terrestrial plants under elevated CO2

A key part of the uncertainty in terrestrial feedbacks on climate change is related to how and to what extent nitrogen (N) availability constrains the stimulation of terrestrial productivity by elevated CO₂ (eCO₂), and whether or not this constraint will become stronger over time. We explored the ecosystem‐scale relationship between responses of plant productivity and N acquisition to eCO₂ in free‐air CO₂ enrichment (FACE) experiments in grassland, cropland and forest ecosystems and found that: (i) in all three ecosystem types, this relationship was positive, linear and strong (r² = 0.68), but exhibited a negative intercept such that plant N acquisition was decreased by 10% when eCO₂ caused neutral or modest changes in productivity. As the ecosystems were markedly N limited, plants with minimal productivity responses to eCO₂ likely acquired less N than ambient CO₂‐grown counterparts because access was decreased, and not because demand was lower. (ii) Plant N concentration was lower under eCO₂, and this decrease was independent of the presence or magnitude of eCO₂‐induced productivity enhancement, refuting the long‐held hypothesis that this effect results from growth dilution. (iii) Effects of eCO₂ on productivity and N acquisition did not diminish over time, while the typical eCO₂‐induced decrease in plant N concentration did. Our results suggest that, at the decennial timescale covered by FACE studies, N limitation of eCO₂‐induced terrestrial productivity enhancement is associated with negative effects of eCO₂ on plant N acquisition rather than with growth dilution of plant N or processes leading to progressive N limitation.     

Forest fine-root production and nitrogen use under elevated CO2: contrasting responses in evergreen and deciduous trees explained by a common principle

Despite the importance of nitrogen (N) limitation of forest carbon (C) sequestration at rising atmospheric CO₂ concentration, the mechanisms responsible are not well understood. To elucidate the interactive effects of elevated CO₂ (eCO₂) and soil N availability on forest productivity and C allocation, we hypothesized that (1) trees maximize fitness by allocating N and C to maximize their net growth and (2) that N uptake is controlled by soil N availability and root exploration for soil N. We tested this model using data collected in Free‐Air CO₂ Enrichment sites dominated by evergreen (Pinus taeda; Duke Forest) and deciduous [Liquidambar styraciflua; Oak Ridge National Laboratory (ORNL)] trees. The model explained 80–95% of variation in productivity and N‐uptake data among eCO₂, N fertilization and control treatments over 6 years. The model explains why fine‐root production increased, and why N uptake increased despite reduced soil N availability under eCO₂ at ORNL and Duke. In agreement with observations at other sites, the model predicts that soil N availability reduced below a critical level diminishes all eCO₂ responses. At Duke, a negative feedback between reduced soil N availability and N uptake prevented progressive reduction in soil N availability at eCO₂. At ORNL, soil N availability progressively decreased because it did not trigger reductions in N uptake; N uptake was maintained at ORNL through a large increase in the production of fast turnover fine roots. This implies that species with fast root turnover could be more prone to progressive N limitation of carbon sequestration in woody biomass than species with slow root turnover, such as evergreens. However, longer term data are necessary for a thorough evaluation of this hypothesis. The success of the model suggests that the principle of maximization of net growth to control growth and allocation could serve as a basis for simplification and generalization of larger scale forest and ecosystem models, for example by removing the need to specify parameters for relative foliage/stem/root allocation.     

Spatial heterogeneity in soil nutrient supply modulates nutrient and biomass responses to multiple global change drivers in model grassland communities

Changes in the atmospheric concentration of carbon dioxide ([CO₂]), nutrient availability and biotic diversity are three major drivers of the ongoing global change impacting terrestrial ecosystems worldwide. While it is well established that soil nutrient heterogeneity exerts a strong influence on the development of plant individuals and communities, it is virtually unknown how nutrient heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. We conducted a microcosm experiment to evaluate the effect of simultaneous changes in [CO₂], nutrient heterogeneity (NH), nutrient availability (NA) and species evenness on the biomass and nutrient uptake patterns of assemblages formed by Lolium perenne, Plantago lanceolata and Holcus lanatus. When the nutrients were heterogeneously supplied, assemblages exhibited precise root foraging patterns, and had higher above‐ and belowground biomass (average increases of 32% and 29% for above‐ and belowground biomass, respectively). Nutrient heterogeneity also modulated the effects of NA on biomass production, complementarity in nitrogen uptake and below: aboveground ratio, as well as those of [CO₂] on the nutrient use efficiency at the assemblage level. Our results show that nutrient heterogeneity has the potential to influence the response of plant assemblages to simultaneous changes in [CO₂], nutrient availability and biotic diversity, and suggest that it is an important environmental factor to interpret and assess plant assemblage responses to global change.     

Biomass production of the last remaining fen with Saxifraga hirculus in Switzerland is controlled by nitrogen availability

Olde Venterink H. and Vittoz P. 2008. Biomass production of the last remaining fen with Saxifraga hirculus in Switzerland is controlled by nitrogen availability. Bot. Helv. 118: 165 – 174. For conservation management of endangered plants it is important to know which nutrient(s) control growth of the vegetation, because maintenance of low nitrogen (N), phosphorus (P) or potassium (K) availability requires different management measures. The aim of this study was to determine the type of nutrient limitation for the vegetation in the last remaining site with Saxifraga hirculus in Switzerland, using nutrient ratios in the aboveground vegetation as an indicator. We made vegetation relevees, collected biomass of the vascular plants, and took soil samples in three plots at this site. The biomass was very low (152–231 g m ^-2), and all three plots were clearly N-limited with N:P ratios of 7– 8. Soil extractable N concentrations were generally low, and P and K concentrations were moderate to high, which was consistent with the indicated N limitation. Hence conservation management first of all needs to prevent N-enrichment, and needs to avoid increased mineralization rates through drainage, or the accumulation of N in the system from atmospheric deposition. Therefore N output seems required through for instance grazing or mowing. The current grazing management seems to function well, since total aboveground biomass is very low and S. hirculus has a high abundance in this last remnant. Submitted 5 June 2008; Accepted 14 October 2008 Subject editor: Sonja Wipf     

Biomass increases attributed to both faster tree growth and altered allometric relationships under long‐term carbon dioxide enrichment at a temperate forest

Increases in atmospheric carbon dioxide (CO₂) concentrations are expected to lead to increases in the rate of tree biomass accumulation, at least temporarily. On the one hand, trees may simply grow faster under higher CO₂ concentrations, preserving the allometric relations that prevailed under lower CO₂ concentrations. Alternatively, the allometric relations themselves may change. In this study, the effects of elevated CO₂ (eCO₂) on tree biomass and allometric relations were jointly assessed. Over 100 trees, grown at Duke Forest, NC, USA, were harvested from eight plots. Half of the plots had been subjected to CO₂ enrichment from 1996 to 2010. Several subplots had also been subjected to nitrogen fertilization from 2005 to 2010. Allometric equations were developed to predict tree height, stem volume, and aboveground biomass components for loblolly pine (Pinus taeda L.), the dominant tree species, and broad‐leaved species. Using the same diameter‐based allometric equations for biomass, it was estimated that plots with eCO₂ contained 21% more aboveground biomass, consistent with previous studies. However, eCO₂ significantly affected allometry, and these changes had an additional effect on biomass. In particular, P. taeda trees at a given diameter were observed to be taller under eCO₂ than under ambient CO₂ due to changes in both the allometric scaling exponent and intercept. Accounting for allometric change increased the treatment effect of eCO₂ on aboveground biomass from a 21% to a 27% increase. No allometric changes for the nondominant broad‐leaved species were identified, nor were allometric changes associated with nitrogen fertilization. For P. taeda, it is concluded that eCO₂ affects allometries, and that knowledge of allometry changes is necessary to accurately compute biomass under eCO₂. Further observations are needed to determine whether this assessment holds for other taxa.     

Response of maize biomass and soil water fluxes on elevated CO2 and drought—From field experiments to process‐based simulations

The rising concentration of atmospheric carbon dioxide (CO₂) is known to increase the total aboveground biomass of several C3 crops, whereas C4 crops are reported to be hardly affected when water supply is sufficient. However, a free‐air carbon enrichment (FACE) experiment in Braunschweig, Germany, in 2007 and 2008 resulted in a 25% increased biomass of the C4 crop maize under restricted water conditions and elevated CO₂ (550 ppm). To project future yields of maize under climate change, an accurate representation of the effects of eCO₂ and drought on biomass and soil water conditions is essential. Current crop growth models reveal limitations in simulations of maize biomass under eCO₂ and limited water supply. We use the coupled process‐based hydrological‐plant growth model Catchment Modeling Framework‐Plant growth Modeling Framework to overcome this limitation. We apply the coupled model to the maize‐based FACE experiment in Braunschweig that provides robust data for the investigation of combined CO₂ and drought effects. We approve hypothesis I that CO₂ enrichment has a small direct‐fertilizing effect with regard to the total aboveground biomass of maize and hypothesis II that CO₂ enrichment decreases water stress and leads to higher yields of maize under restricted water conditions. Hypothesis III could partly be approved showing that CO₂ enrichment decreases the transpiration of maize, but does not raise soil moisture, while increasing evaporation. We emphasize the importance of plant‐specific CO₂ response factors derived by use of comprehensive FACE data. By now, only one FACE experiment on maize is accomplished applying different water levels. For the rigorous testing of plant growth models and their applicability in climate change studies, we call for datasets that go beyond single criteria (only yield response) and single effects (only elevated CO₂).     

Elevated atmospheric carbon dioxide impairs the performance of root‐feeding vine weevils by modifying root growth and secondary metabolites

Predicting how insect crop pests will respond to global climate change is an important part of increasing crop production for future food security, and will increasingly rely on empirically based evidence. The effects of atmospheric composition, especially elevated carbon dioxide (eCO₂), on insect herbivores have been well studied, but this research has focussed almost exclusively on aboveground insects. However, responses of root‐feeding insects to eCO₂ are unlikely to mirror these trends because of fundamental differences between aboveground and belowground habitats. Moreover, changes in secondary metabolites and defensive responses to insect attack under eCO₂ conditions are largely unexplored for root herbivore interactions. This study investigated how eCO₂ (700 μmol mol^?1) affected a root‐feeding herbivore via changes to plant growth and concentrations of carbon (C), nitrogen (N) and phenolics. This study used the root‐feeding vine weevil, Otiorhynchus sulcatus and the perennial crop, Ribes nigrum. Weevil populations decreased by 33% and body mass decreased by 23% (from 7.2 to 5.4 mg) in eCO₂. Root biomass decreased by 16% in eCO₂, which was strongly correlated with weevil performance. While root N concentrations fell by 8%, there were no significant effects of eCO₂ on root C and N concentrations. Weevils caused a sink in plants, resulting in 8–12% decreases in leaf C concentration following herbivory. There was an interactive effect of CO₂ and root herbivory on root phenolic concentrations, whereby weevils induced an increase at ambient CO₂, suggestive of defensive response, but caused a decrease under eCO₂. Contrary to predictions, there was a positive relationship between root phenolics and weevil performance. We conclude that impaired root‐growth underpinned the negative effects of eCO₂ on vine weevils and speculate that the plant's failure to mount a defensive response at eCO₂ may have intensified these negative effects.     

Impacts of long‐term elevated atmospheric CO2 concentrations on communities of arbuscular mycorrhizal fungi

The ecological impacts of long‐term elevated atmospheric CO₂ (eCO₂) levels on soil microbiota remain largely unknown. This is particularly true for the arbuscular mycorrhizal (AM) fungi, which form mutualistic associations with over two‐thirds of terrestrial plant species and are entirely dependent on their plant hosts for carbon. Here, we use high‐resolution amplicon sequencing (Illumina, HiSeq) to quantify the response of AM fungal communities to the longest running (>15 years) free‐air carbon dioxide enrichment (FACE) experiment in the Northern Hemisphere (GiFACE); providing the first evaluation of these responses from old‐growth (>100 years) semi‐natural grasslands subjected to a 20% increase in atmospheric CO₂. eCO₂ significantly increased AM fungal richness but had a less‐pronounced impact on the composition of their communities. However, while broader changes in community composition were not observed, more subtle responses of specific AM fungal taxa were with populations both increasing and decreasing in abundance in response to eCO₂. Most population‐level responses to eCO₂ were not consistent through time, with a significant interaction between sampling time and eCO₂ treatment being observed. This suggests that the temporal dynamics of AM fungal populations may be disturbed by anthropogenic stressors. As AM fungi are functionally differentiated, with different taxa providing different benefits to host plants, changes in population densities in response to eCO₂ may significantly impact terrestrial plant communities and their productivity. Thus, predictions regarding future terrestrial ecosystems must consider changes both aboveground and belowground, but avoid relying on broad‐scale community‐level responses of soil microbes observed on single occasions.     

Biomass responses in a temperate European grassland through 17 years of elevated CO2

Future increase in atmospheric CO₂ concentrations will potentially enhance grassland biomass production and shift the functional group composition with consequences for ecosystem functioning. In the “GiFACE” experiment (Giessen Free Air Carbon dioxide Enrichment), fertilized grassland plots were fumigated with elevated CO₂ (eCO₂) year‐round during daylight hours since 1998, at a level of +20% relative to ambient concentrations (in 1998, aCO₂ was 364 ppm and eCO₂ 399 ppm; in 2014, aCO₂ was 397 ppm and eCO₂ 518 ppm). Harvests were conducted twice annually through 23 years including 17 years with eCO₂ (1998 to 2014). Biomass consisted of C3 grasses and forbs, with a small proportion of legumes. The total aboveground biomass (TAB) was significantly increased under eCO₂ (p = .045 and .025, at first and second harvest). The dominant plant functional group grasses responded positively at the start, but for forbs, the effect of eCO₂ started out as a negative response. The increase in TAB in response to eCO₂ was approximately 15% during the period from 2006 to 2014, suggesting that there was no attenuation of eCO₂ effects over time, tentatively a consequence of the fertilization management. Biomass and soil moisture responses were closely linked. The soil moisture surplus (c. 3%) in eCO₂ manifested in the latter years was associated with a positive biomass response of both functional groups. The direction of the biomass response of the functional group forbs changed over the experimental duration, intensified by extreme weather conditions, pointing to the need of long‐term field studies for obtaining reliable responses of perennial ecosystems to eCO₂ and as a basis for model development.     

Strong photosynthetic acclimation and enhanced water‐use efficiency in grassland functional groups persist over 21 years of CO2 enrichment,independent of nitrogen supply

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.     

Influence of experimental,environmental, and geographic factors on nutrient‐diffusing substrate experiments in running waters

相似文献   

Phosphorus status determines biomass response to elevated CO2 in a legume : C4 grass community

Thirty-six mesocosms, each containing a two-species community of Trifolium repens (C₃ legume) and Stenotaphrum secundatum (C₄ grass), were grown in sand with three nutrient regimes, zero N low P, zero N high P and supplied N high P, under ambient (aCO₂) and twice ambient CO₂ (eCO₂) for 15 months in two greenhouses. Aboveground annual production in the P limited mesocosms did not respond to eCO₂ and was reduced by 50% relative to mesocosms with an adequate P supply, where dry-matter production was increased by 12–24% under eCO₂. The stimulation of production by eCO₂ occurred throughout the year despite a clear seasonality in growth. There was no effect of eCO₂ on leaf area index (LAI), which was larger under high P than low P. Live root mass at the end of the experiment was higher under eCO₂ in all nutrient treatments, but the response of total belowground C (root+soil) to eCO₂ depended on P treatment. Under limiting P, belowground C was not significantly changed by eCO₂ (2–2.3 t belowground C ha⁻¹). Under high P supply, both root and soil C pools increased under eCO₂. Under aCO₂, low P supply increased belowground C by 0.7–1 t C ha⁻¹ above that added by the high P treatment. P is commonly limiting in Australian ecosystems and the majority of ecosystem N input is provided by biological N fixation. Consequently, the response of legumes to eCO₂ is of particular importance. These results demonstrate that at low P availability, there is likely to be only a limited response of biomass production by T. repens to eCO₂, which in turn may constrain any ecosystem response.     

Alteration of forest succession and carbon cycling under elevated CO2

Regenerating forests influence the global carbon (C) cycle, and understanding how climate change will affect patterns of regeneration and C storage is necessary to predict the rate of atmospheric carbon dioxide (CO₂) increase in future decades. While experimental elevation of CO₂ has revealed that young forests respond with increased productivity, there remains considerable uncertainty as to how the long‐term dynamics of forest regrowth are shaped by elevated CO₂ (eCO₂). Here, we use the mechanistic size‐ and age‐ structured Ecosystem Demography model to investigate the effects of CO₂ enrichment on forest regeneration, using data from the Duke Forest Free‐Air Carbon dioxide Enrichment (FACE) experiment, a forest chronosequence, and an eddy‐covariance tower for model parameterization and evaluation. We find that the dynamics of forest regeneration are accelerated, and stands consistently hit a variety of developmental benchmarks earlier under eCO₂. Because responses to eCO₂ varied by plant functional type, successional pathways, and mature forest composition differed under eCO₂, with mid‐ and late‐successional hardwood functional types experiencing greater increases in biomass compared to early‐successional functional types and the pine canopy. Over the simulation period, eCO₂ led to an increase in total ecosystem C storage of 9.7 Mg C ha^‐1. Model predictions of mature forest biomass and ecosystem–atmosphere exchange of CO₂ and H₂O were sensitive to assumptions about nitrogen limitation; both the magnitude and persistence of the ecosystem response to eCO₂ were reduced under N limitation. In summary, our simulations demonstrate that eCO₂ can result in a general acceleration of forest regeneration while altering the course of successional change and having a lasting impact on forest ecosystems.