Combined effects of elevated CO2 concentration and Wolbachia on Hylyphantes graminicola (Araneae: Linyphiidae)

The increasing concentration of carbon dioxide in atmosphere is not only a major cause of global warming, but it also adversely affects the ecological diversity of invertebrates. This study was conducted to evaluate the effect of elevated CO₂ concentration (ambient, 400 ppm and high, 800 ppm) and Wolbachia (Wolbachia‐infected, W⁺ and Wolbachia‐uninfected, W^?) on Hylyphantes graminicola. The total survival rate, developmental duration, carapace width and length, body weight, sex ratio, net reproductive rate, nutrition content, and enzyme activity in H. graminicola were examined under four treatments: W^? 400 ppm, W^? 800 ppm, W⁺ 400 ppm, and W⁺ 800 ppm. Results showed that Wolbachia‐infected spiders had significantly decreased the total developmental duration. Different instars showed variations up to some extent, but no obvious effect was found under elevated CO₂ concentration. Total survival rate, sex ratio, and net reproductive rate were not affected by elevated CO₂ concentration or Wolbachia infection. The carapace width of Wolbachia‐uninfected spiders decreased significantly under elevated CO₂ concentration, while the width, length and weight were not significantly affected in Wolbachia‐infected spiders reared at ambient CO₂ concentration. The levels of protein, specific activities of peroxidase, and amylase were significantly increased under elevated CO₂ concentration or Wolbachia‐infected spiders, while the total amino content was only increased in Wolbachia‐infected spiders. Thus, our current finding suggested that elevated CO₂ concentration and Wolbachia enhance nutrient contents and enzyme activity of H. graminicola and decrease development duration hence explore the interactive effects of factors which were responsible for reproduction regulation, but it also gives a theoretical direction for spider's protection in such a dynamic environment. Increased activities of enzymes and nutrients caused by Wolbachia infection aids for better survival of H. graminicola under stress.     

The effect of atmospheric carbon dioxide concentrations on the performance of the mangrove Avicennia germinans over a range of salinities

By increasing water use efficiency and carbon assimilation, increasing atmospheric CO₂ concentrations could potentially improve plant productivity and growth at high salinities. To assess the effect of elevated CO₂ on the salinity response of a woody halophyte, we grew seedlings of the mangrove Avicennia germinans under a combination of five salinity treatments [from 5 to 65 parts per thousand (ppt)] and three CO₂ concentrations (280, 400 and 800 ppm). We measured survivorship, growth rate, photosynthetic gas exchange, root architecture and foliar nutrient and ion concentrations. The salinity optima for growth shifted higher with increasing concentrations of CO₂, from 0 ppt at 280 ppm to 35 ppt at 800 ppm. At optimal salinity conditions, carbon assimilation rates were significantly higher under elevated CO₂ concentrations. However, at salinities above the salinity optima, salinity had an expected negative effect on mangrove growth and carbon assimilation, which was not alleviated by elevated CO₂, despite a significant improvement in photosynthetic water use efficiency. This is likely due to non‐stomatal limitations to growth at high salinities, as indicated by our measurements of foliar ion concentrations that show a displacement of K⁺ by Na⁺ at elevated salinities that is not affected by CO₂. The observed shift in the optimal salinity for growth with increasing CO₂ concentrations changes the fundamental niche of this species and could have significant effects on future mangrove distribution patterns and interspecific interactions.     

Elevated CO2 plus chronic warming reduce nitrogen uptake and levels or activities of nitrogen‐uptake and ‐assimilatory proteins in tomato roots

Atmospheric CO₂ enrichment is expected to often benefit plant growth, despite causing global warming and nitrogen (N) dilution in plants. Most plants primarily procure N as inorganic nitrate (NO₃^?) or ammonium (NH₄⁺), using membrane‐localized transport proteins in roots, which are key targets for improving N use. Although interactive effects of elevated CO₂, chronic warming and N form on N relations are expected, these have not been studied. In this study, tomato (Solanum lycopersicum) plants were grown at two levels of CO₂ (400 or 700 ppm) and two temperature regimes (30 or 37°C), with NO₃^? or NH₄⁺ as the N source. Elevated CO₂ plus chronic warming severely inhibited plant growth, regardless of N form, while individually they had smaller effects on growth. Although %N in roots was similar among all treatments, elevated CO₂ plus warming decreased (1) N‐uptake rate by roots, (2) total protein concentration in roots, indicating an inhibition of N assimilation and (3) shoot %N, indicating a potential inhibition of N translocation from roots to shoots. Under elevated CO₂ plus warming, reduced NO₃^?‐uptake rate per g root was correlated with a decrease in the concentration of NO₃^?‐uptake proteins per g root, reduced NH₄⁺ uptake was correlated with decreased activity of NH₄⁺‐uptake proteins and reduced N assimilation was correlated with decreased concentration of N‐assimilatory proteins. These results indicate that elevated CO₂ and chronic warming can act synergistically to decrease plant N uptake and assimilation; hence, future global warming may decrease both plant growth and food quality (%N).     

Photosynthetic downregulation in leaves of the Japanese white birch grown under elevated CO2 concentration does not change their temperature‐dependent susceptibility to photoinhibition

To determine the effects of elevated CO₂ concentration ([CO₂]) on the temperature‐dependent photosynthetic properties, we measured gas exchange and chlorophyll fluorescence at various leaf temperatures (15, 20, 25, 30, 35 and 40°C) in 1‐year‐old seedlings of the Japanese white birch (Betula platyphylla var. japonica), grown in a phytotron under natural daylight at two [CO₂] levels (ambient: 400 µmol mol^?1 and elevated: 800 µmol mol^?1) and limited N availability (90 mg N plant^?1). Plants grown under elevated [CO₂] exhibited photosynthetic downregulation, indicated by a decrease in the carboxylation capacity of Rubisco. At temperatures above 30°C, the net photosynthetic rates of elevated‐CO₂‐grown plants exceeded those grown under ambient [CO₂] when compared at their growth [CO₂]. Electron transport rates were significantly lower in elevated‐CO₂‐grown plants than ambient‐CO₂‐grown ones at temperatures below 25°C. However, no significant difference was observed in the fraction of excess light energy [(1 ? q_P)× F_v′/F_m′] between CO₂ treatments across the temperature range. The quantum yield of regulated non‐photochemical energy loss was significantly higher in elevated‐CO₂‐grown plants than ambient, when compared at their respective growth [CO₂] below 25°C. These results suggest that elevated‐CO₂‐induced downregulation might not exacerbate the temperature‐dependent susceptibility to photoinhibition, because reduced energy consumption by electron transport was compensated for by increased thermal energy dissipation at low temperatures.     

Interspecies variation in nitrogen uptake kinetic responses of temperate forest species to elevated CO2: potential causes and consequences

Despite the recognition that the capacity to acquire N is critical in plant response to CO₂ enrichment, there is little information on how elevated CO₂ affects root N uptake kinetics. The few available data indicate a highly variable pattern of response to elevated CO₂, but it is presently unclear if the observed inconsistencies are caused by differences in experimental protocols or by true species differences. Furthermore, if there are interspecific variations in N uptake responses to elevated CO₂, it is not clear whether these are associated with different functional groups. Accordingly, we examined intact root‐system NH₄⁺ and NO₃^– uptake kinetic responses to elevated CO₂ in seedlings of six temperate forest tree species, representing (i) fast‐ vs. slow‐growers and (ii) broad‐leaves vs. conifers, that were cultured and assayed in otherwise similar conditions. In general, the species tested had a higher uptake capacity (V_max) for NH₄⁺ than for NO₃^–. Species substantially differed in their NO₃^– and NH₄⁺ uptake capacities, but the interspecific differences were markedly greater for NO₃^– than NH₄⁺ uptake. Elevated CO₂ had a species‐dependent effect on root uptake capacity for NH₄⁺ ranging from an increase of 215% in Acer negundo L. to a decrease of about 40% in Quercus macrocarpa Michx. In contrast, NO₃^– uptake capacity responded little to CO₂ in all the species except A. negundo in which it was significantly down‐regulated at elevated CO₂. Across species, the capacity for NH₄⁺ uptake was positively correlated with the relative growth rate (RGR) of species; however, the CO₂ effect on NH₄⁺ uptake capacity could not be explained by changes in RGR. The observed variation in NH₄⁺ uptake response to elevated CO₂ was also inconsistent with life‐form differences. Other possible mechanisms that may explain why elevated CO₂ elicits a species‐specific response in root N uptake kinetics are discussed. Despite the fact that the exact mechanism(s) for such interspecific variation remains unresolved, these differences may have a significant implication for competitive interactions and community responses to elevated CO₂ environment. We suggest that differential species responses in nutrient uptake capacity could be one potential mechanism for the CO₂‐induced shifts in net primary productivity and species composition that have been observed in experimental communities exposed to elevated levels of CO₂.     

A cycad's non‐saturating response to carbon dioxide enrichment indicates Cenozoic carbon limitation in pre‐historic plants

Cycads were a dominant plant functional type during the Mesozoic Era when atmospheric carbon dioxide [CO₂] greatly exceeded current conditions. Cycads, now rare and endangered, are slow‐growing perennial gymnosperms that develop carbon‐rich structural biomass, such as sclerophyllous leaves, dense stems and massive reproductive cones. Is cycad carbon partitioning to specific organs a constraint of their high [CO₂] evolutionary history (CO₂ legacy hypothesis, CLH)? To explore changes in cycad growth, carbon partitioning and assimilation responses that could be expected during the CO₂ depletion of the Cenozoic Era, individuals of the cycad species Encephalartos villosus plants were grown at four CO₂ levels: 400, 550, 750 and 1000 μmol mol^?1. The CLH predicts that cycad biomass and growth rates would increase in elevated [CO₂] due to increased net assimilation rates, and that carbon‐dense structures would provide sufficient carbohydrate sinks to prevent photosynthetic down‐regulation even under super‐ambient [CO₂] of 1000 μmol mol^?1. Both hypotheses were confirmed, though the latter less strongly. Plant relative growth rates increased 23% and biomass accumulation increased 65% in 1000 μmol mol^?1relative to 400 μmol mol^?1 treatment groups. Mean net assimilation rates increased 130% at 1000 μmol mol^?1 relative to 400 μmol mol^?1 CO₂, though there was some down‐regulation of maximum rate of carboxylation (Vc_max). Assimilation rates, relative growth rates, biomass and mean leaf sugar content were linearly related to [CO₂] over the entire experimental range. Photosynthesis appears to be regulated by stomata at low CO₂ levels and by non‐stomatal (i.e. biochemical limitations) at greater concentrations. In general, our results suggest that growth and physiological performance of cycads have been severely compromised by declining [CO₂] during the Cenozoic Era, possibly contributing to the current rare and endangered status of this functional type.     

Nutrient supply enhanced the increase in intrinsic water‐use efficiency of a temperate seminatural grassland in the last century

Under the increase in atmospheric CO₂ during the last century, variable increases in the intrinsic water‐use efficiency (W_i), i.e., the ratio between carbon assimilation rate (A) and stomatal conductance (g_s), of C₃ vegetation have been observed. Here, we ask if long‐term nutrient status and especially nitrogen supply have an effect on the CO₂ response of W_i in a temperate seminatural C₃ grassland. This analysis draws on the long‐term trends (1915–2009) in W_i, derived from carbon isotope analysis, of archived hay and herbage from the Park Grass Experiment at Rothamsted (South‐East England). Plant samples came from five fertilizer treatments, each with different annual nitrogen (N; 0, 48 or 96 kg ha^?1), phosphorus (P; 0 or 35 kg ha^?1) and potassium (K; 0 or 225 kg ha^?1) applications, with lime as required to maintain soil pH near 7. Carbon isotope discrimination (¹³Δ) increased significantly (P < 0.001) on the Control (0.9‰ per 100 ppm CO₂ increase). This trend differed significantly (P < 0.01) from those observed on the fertilized treatments (PK only: 0.4‰ per 100 ppm CO₂ increase, P < 0.001; Low N only, Low N+PK, High N+PK: no significant increase). The ¹³Δ trends on fertilized treatments did not differ significantly from each other. However, N status, assessed as N fertilizer supply plus an estimate of biologically fixed N, was negatively related (r² = 0.88; P < 0.02) to the trend for ¹³Δ against CO₂. Other indices of N status exhibited similar relationships. Accordingly, the increase in W_i at High N+PK was twice that of the Control (+28% resp. +13% relative to 1915). In addition, the CO₂ responsiveness of ¹³Δ was related to the grass content of the plant community. This may have been due to the greater CO₂ responsiveness of g_s in grasses relative to forbs. Thus, the greater CO₂ response of grass‐rich fertilized swards may be related to effects of nutrient supply on botanical composition.     

Male rescue maintains low frequency parthenogenesis-inducing <Emphasis Type="Italic">Wolbachia</Emphasis> infection in <Emphasis Type="Italic">Trichogramma</Emphasis> populations

Parthenogenesis-inducing (PI) Wolbachia bacteria are reproductive parasites that cause infected (W ⁺) female haplodiploid parasitoids to produce daughters without fertilization by males. Theoretically, PI Wolbachia infection should spread to fixation within Trichogramma populations as males are no longer required to produce female offspring. Infections in some naturally occurring Trichogramma populations are, however, maintained at frequencies ranging from 4 to 26%. Here we describe discrete equation models to examine if the PI Wolbachia infection in Trichogramma populations can be maintained at relatively low frequencies by mating regularity. Model outcomes suggest the probability of W ⁺ females mating could stabilize Wolbachia infection frequency at low levels in Trichogramma populations. The primary mechanism maintaining low-level PI Wolbachia infection in Trichogramma populations is reducing the survivorship from egg to adult in infected relative to uninfected females. The model successfully demonstrates that the relatively low PI Wolbachia infection frequency in host populations can be maintained by fertilization, or male rescue, of infected eggs, which avoids potentially hazardous gamete duplication that occurs during Wolbachia-induced parthenogenesis.     

Virus disease in wheat predicted to increase with a changing climate

Current atmospheric CO₂ levels are about 400 μmol mol^?1 and are predicted to rise to 650 μmol mol^?1 later this century. Although the positive and negative impacts of CO₂ on plants are well documented, little is known about interactions with pests and diseases. If disease severity increases under future environmental conditions, then it becomes imperative to understand the impacts of pathogens on crop production in order to minimize crop losses and maximize food production. Barley yellow dwarf virus (BYDV) adversely affects the yield and quality of economically important crops including wheat, barley and oats. It is transmitted by numerous aphid species and causes a serious disease of cereal crops worldwide. This study examined the effects of ambient (aCO₂; 400 μmol mol^?1) and elevated CO₂ (eCO₂; 650 μmol mol^?1) on noninfected and BYDV‐infected wheat. Using a RT‐qPCR technique, we measured virus titre from aCO₂ and eCO₂ treatments. BYDV titre increased significantly by 36.8% in leaves of wheat grown under eCO₂ conditions compared to aCO₂. Plant growth parameters including height, tiller number, leaf area and biomass were generally higher in plants exposed to higher CO₂ levels but increased growth did not explain the increase in BYDV titre in these plants. High virus titre in plants has been shown to have a significant negative effect on plant yield and causes earlier and more pronounced symptom expression increasing the probability of virus spread by insects. The combination of these factors could negatively impact food production in Australia and worldwide under future climate conditions. This is the first quantitative evidence that BYDV titre increases in plants grown under elevated CO₂ levels.     

Effects of elevated carbon dioxide on stomatal characteristics and carbon isotope ratio of Arabidopsis thaliana ecotypes originating from an altitudinal gradient

Stomatal functioning regulates the fluxes of CO₂ and water vapor between vegetation and atmosphere and thereby influences plant adaptation to their habitats. Stomatal traits are controlled by external environmental and internal cellular signaling. The objective of this study was to quantify the effects of CO₂ enrichment (CE) on stomatal density (SD)‐related properties, guard cell length (GCL) and carbon isotope ratio (δ¹³C) of a range of Arabidopsis thaliana ecotypes originating from a wide altitudinal range [50–1260 m above sea level (asl)], and grown at 400 and 800 ppm [CO₂], and thereby elucidate the possible adaptation and acclimation responses controlling stomatal traits and water use efficiency (WUE). There was a highly significant variation among ecotypes in the magnitude and direction of response of stomatal traits namely, SD and stomatal index (SI) and GCL, and δ¹³C to CE, which represented a short‐term acclimation response. A majority of ecotypes showed increased SD and SI with CE with the response not depending on the altitude of origin. Significant ecotypic variation was shown in all stomatal traits and δ¹³C at each [CO₂]. At 400 ppm, means of SD, SI and GCL for broad altitudinal ranges, i.e. low (<100 m), mid (100–400 m) and high (>400 m), increased with increasing altitude, which represented an adaptation response to decreased availability of CO₂ with altitude. δ¹³C was negatively correlated to SD and SI at 800 ppm but not at 400 ppm. Our results highlight the diversity in the response of key stomatal characters to CE and altitude within the germplasm of A. thaliana and the need to consider this diversity when using A. thaliana as a model plant.     

Interactive direct and plant‐mediated effects of elevated atmospheric [CO2] and temperature on a eucalypt‐feeding insect herbivore

Understanding the direct and indirect effects of elevated [CO₂] and temperature on insect herbivores and how these factors interact are essential to predict ecosystem‐level responses to climate change scenarios. In three concurrent glasshouse experiments, we measured both the individual and interactive effects of elevated [CO₂] and temperature on foliar quality. We also assessed the interactions between their direct and plant‐mediated effects on the development of an insect herbivore of eucalypts. Eucalyptus tereticornis saplings were grown at ambient or elevated [CO₂] (400 and 650 μmol mol^?1 respectively) and ambient or elevated ( + 4 °C) temperature for 10 months. Doratifera quadriguttata (Lepidoptera: Limacodidae) larvae were feeding directly on these trees, on their excised leaves in a separate glasshouse, or on excised field‐grown leaves within the temperature and [CO₂] controlled glasshouse. To allow insect gender to be determined and to ensure that any sex‐specific developmental differences could be distinguished from treatment effects, insect development time and consumption were measured from egg hatch to pupation. No direct [CO₂] effects on insects were observed. Elevated temperature accelerated larval development, but did not affect leaf consumption. Elevated [CO₂] and temperature independently reduced foliar quality, slowing larval development and increasing consumption. Simultaneously increasing both [CO₂] and temperature reduced these shifts in foliar quality, and negative effects on larval performance were subsequently ameliorated. Negative nutritional effects of elevated [CO₂] and temperature were also independently outweighed by the direct positive effect of elevated temperature on larvae. Rising [CO₂] and temperature are thus predicted to have interactive effects on foliar quality that affect eucalypt‐feeding insects. However, the ecological consequences of these interactions will depend on the magnitude of concurrent temperature rise and its direct effects on insect physiology and feeding behaviour.     

Rising CO2 from historical concentrations enhances the physiological performance of Brassica napus seedlings under optimal water supply but not under reduced water availability

The productivity of many important crops is significantly threatened by water shortage, and the elevated atmospheric CO₂ can significantly interact with physiological processes and crop responses to drought. We examined the effects of three different CO₂ concentrations (historical ~300 ppm, ambient ~400 ppm and elevated ~700 ppm) on physiological traits of oilseed rape (Brassica napus L.) seedlings subjected to well‐watered and reduced water availability. Our data show (1) that, as expected, increasing CO₂ level positively modulates leaf photosynthetic traits, leaf water‐use efficiency and growth under non‐stressed conditions, although a pronounced acclimation of photosynthesis to elevated CO₂ occurred; (2) that the predicted elevated CO₂ concentration does not reduce total evapotranspiration under drought when compared with present (400 ppm) and historical (300 ppm) concentrations because of a larger leaf area that does not buffer transpiration; and (3) that accordingly, the physiological traits analysed decreased similarly under stress for all CO₂ concentrations. Our data support the hypothesis that increasing CO₂ concentrations may not significantly counteract the negative effect of increasing drought intensity on Brassica napus performance.     

Population parameters and growth of Riptortus pedestris (Fabricius) (Hemiptera: Alydidae) under elevated CO2 concentrations

We evaluated the direct effects of three different CO₂ concentrations (400, 600 and 1,000 ppm) on the population parameters and growth of the bean bug, Riptortus pedestris, while being fed on soybean. The raw life history data from R. pedestris was analyzed using an age‐stage, two‐sex life table to take the viable development rate among individuals into account. Based on the age‐stage, two‐sex life table analysis, the population projections of R. pedestris provide the stage structure and variability of the population growth under different CO₂ treatments. Our results showed significantly shorter immature durations and higher pre‐adult survival rate under elevated CO₂ (1,000 ppm) than those under ambient CO₂ (400 ppm). The population of R. pedestris reared under elevated CO₂ conditions showed higher intrinsic and finite rates of increase but a lower mean generation time than R. pedestris reared under ambient CO₂ conditions. Our results show the population parameters and growth of R. pedestris are influenced by increased CO₂ relative to ambient CO₂ treatment. Further studies on the long‐term direct effects of different CO₂ levels on R. pedestris are essential to understand their population dynamics and to establish appropriate management strategies.     

Interactive effects of elevated CO2 and soil fertility on isoprene emissions from Quercus robur

The effects of global change on the emission rates of isoprene from plants are not clear. A factor that can influence the response of isoprene emission to elevated CO₂ concentrations is the availability of nutrients. Isoprene emission rate under standard conditions (leaf temperature: 30°C, photosynthetically active radiation (PAR): 1000 μmol photons m^?2 s^?1), photosynthesis, photosynthetic capacity, and leaf nitrogen (N) content were measured in Quercus robur grown in well‐ventilated greenhouses at ambient and elevated CO₂ (ambient plus 300 ppm) and two different soil fertilities. The results show that elevated CO₂ enhanced photosynthesis but leaf respiration rates were not affected by either the CO₂ or nutrient treatments. Isoprene emission rates and photosynthetic capacity were found to decrease with elevated CO₂, but an increase in nutrient availability had the converse effect. Leaf N content was significantly greater with increased nutrient availability, but unaffected by CO₂. Isoprene emission rates measured under these conditions were strongly correlated with photosynthetic capacity across the range of different treatments. This suggests that the effects of CO₂ and nutrient levels on allocation of carbon to isoprene production and emission under near‐saturating light largely depend on the effects on photosynthetic electron transport capacity.     

Direct and indirect effects of elevated atmospheric CO2 on net ecosystem production in a Chesapeake Bay tidal wetland

The rapid increase in atmospheric CO₂ concentrations (C_a) has resulted in extensive research efforts to understand its impact on terrestrial ecosystems, especially carbon balance. Despite these efforts, there are relatively few data comparing net ecosystem exchange of CO₂ between the atmosphere and the biosphere (NEE), under both ambient and elevated C_a. Here we report data on annual sums of CO₂ (NEE_net) for 19 years on a Chesapeake Bay tidal wetland for Scirpus olneyi (C₃ photosynthetic pathway)‐ and Spartina patens (C₄ photosynthetic pathway)‐dominated high marsh communities exposed to ambient and elevated C_a (ambient + 340 ppm). Our objectives were to (i) quantify effects of elevated C_a on seasonally integrated CO₂ assimilation (NEE_net = NEE_day + NEE_night, kg C m^?2 y^?1) for the two communities; and (ii) quantify effects of altered canopy N content on ecosystem photosynthesis and respiration. Across all years, NEE_net averaged 1.9 kg m^?2 y^?1 in ambient C_a and 2.5 kg m^?2 y^?1 in elevated C_a, for the C₃‐dominated community. Similarly, elevated C_a significantly (P < 0.01) increased carbon uptake in the C₄‐dominated community, as NEE_net averaged 1.5 kg m^?2 y^?1 in ambient C_a and 1.7 kg m^?2 y^?1 in elevated C_a. This resulted in an average CO₂ stimulation of 32% and 13% of seasonally integrated NEE_net for the C₃‐ and C₄‐dominated communities, respectively. Increased NEE_day was correlated with increased efficiencies of light and nitrogen use for net carbon assimilation under elevated C_a, while decreased NEE_night was associated with lower canopy nitrogen content. These results suggest that rising C_a may increase carbon assimilation in both C₃‐ and C₄‐dominated wetland communities. The challenge remains to identify the fate of the assimilated carbon.     

Does growth under elevated CO2 moderate photoacclimation in rice?

Acclimation of plant photosynthesis to light irradiance (photoacclimation) involves adjustments in levels of pigments and proteins and larger scale changes in leaf morphology. To investigate the impact of rising atmospheric CO₂ on crop physiology, we hypothesize that elevated CO₂ interacts with photoacclimation in rice (Oryza sativa). Rice was grown under high light (HL: 700 µmol m^?2 s^?1), low light (LL: 200 µmol m^?2 s^?1), ambient CO₂ (400 µl l^?1) and elevated CO₂ (1000 µl l^?1). Leaf six was measured throughout. Obscuring meristem tissue during development did not alter leaf thickness indicating that mature leaves are responsible for sensing light during photoacclimation. Elevated CO₂ raised growth chamber photosynthesis and increased tiller formation at both light levels, while it increased leaf length under LL but not under HL. Elevated CO₂ always resulted in increased leaf growth rate and tiller production. Changes in leaf thickness, leaf area, Rubisco content, stem and leaf starch, sucrose and fructose content were all dominated by irradiance and unaffected by CO₂. However, stomata responded differently; they were significantly smaller in LL grown plants compared to HL but this effect was significantly suppressed under elevated CO₂. Stomatal density was lower under LL, but this required elevated CO₂ and the magnitude was adaxial or abaxial surface‐dependent. We conclude that photoacclimation in rice involves a systemic signal. Furthermore, extra carbohydrate produced under elevated CO₂ is utilized in enhancing leaf and tiller growth and does not enhance or inhibit any feature of photoacclimation with the exception of stomatal morphology.     

Growth of Eastern Cottonwoods (Populus deltoides) in elevated [CO2] stimulates stand-level respiration and rhizodeposition of carbohydrates, accelerates soil nutrient depletion, yet stimulates above- and belowground biomass production

We took advantage of the distinctive system‐level measurement capabilities of the Biosphere 2 Laboratory (B2L) to examine the effects of prolonged exposure to elevated [CO₂] on carbon flux dynamics, above‐ and belowground biomass changes, and soil carbon and nutrient capital in plantation forest stands over 4 years. Annually coppiced stands of eastern cottonwoods (Populus deltoides) were grown under ambient (400 ppm) and two levels of elevated (800 and 1200 ppm) atmospheric [CO₂] in carbon and N‐replete soils of the Intensive Forestry Mesocosm in the B2L. The large semiclosed space of B2L uniquely enabled precise CO₂ exchange measurements at the near ecosystem scale. Highly controllable climatic conditions within B2L also allowed for reproducible examination of CO₂ exchange under different scales in space and time. Elevated [CO₂] significantly stimulated whole‐system maximum net CO₂ influx by an average of 21% and 83% in years 3 and 4 of the experiment. Over the 4‐year experiment, cumulative belowground, foliar, and total aboveground biomass increased in both elevated [CO₂] treatments. After 2 years of growth at elevated [CO₂], early season stand respiration was decoupled from CO₂ influx aboveground, presumably because of accelerated fine root production from stored carbohydrates in the coppiced system prior to canopy development and to the increased soil carbohydrate status under elevated [CO₂] treatments. Soil respiration was stimulated by elevated [CO₂] whether measured at the system level in the undisturbed soil block, by soil collars in situ, or by substrate‐induced respiration in vitro. Elevated [CO₂] accelerated depletion of soil nutrients, phosphorus, calcium and potassium, after 3 years of growth, litter removal, and coppicing, especially in the upper soil profile, although total N showed no change. Enhancement of above‐ and belowground biomass production by elevated [CO₂] accelerated carbon cycling through the coppiced system and did not sequester additional carbon in the soil.     

Soil nitrogen transformations under Populus tremuloides, Betula papyrifera and Acer saccharum following 3 years exposure to elevated CO2 and O3

Increases in atmospheric CO₂ and tropospheric O₃ may affect forest N cycling by altering plant litter production and the availability of substrates for microbial metabolism. Three years following the establishment of our free‐air CO₂–O₃ enrichment experiment, plant growth has been stimulated by elevated CO₂ resulting in greater substrate input to soil; elevated O₃ has counteracted this effect. We hypothesized that rates of soil N cycling would be enhanced by greater plant productivity under elevated CO₂, and that CO₂ effects would be dampened by O₃. We found that elevated CO₂ did not alter gross N transformation rates. Elevated O₃ significantly reduced gross N mineralization and microbial biomass N, and effects were consistent among species. We also observed significant interactions between CO₂ and O₃: (i) gross N mineralization was greater under elevated CO₂ (1.0 mg N kg^?1 day^?1) than in the presence of both CO₂ and O₃ (0.5 mg N kg^?1 day^?1) and (ii) gross NH₄⁺ immobilization was also greater under elevated CO₂ (0.8 mg N kg^?1 day^?1) than under CO₂ plus O₃ (0.4 mg N kg^?1 day^?1). We used a laboratory ¹⁵N tracer method to quantify transfer of inorganic N to organic pools. Elevated CO₂ led to greater recovery of NH₄⁺‐¹⁵N in microbial biomass and corresponding lower recovery in the extractable NO₃^? pool. Elevated CO₂ resulted in a substantial increase in NO₃^?‐¹⁵N recovery in soil organic matter. We observed no O₃ main effect and no CO₂ by O₃ interaction effect on ¹⁵N recovery in any soil pool. All of the above responses were most pronounced beneath Betula papyrifera and Populus tremuloides, which have grown more rapidly than Acer saccharum. Although elevated CO₂ has increased plant productivity, the resulting increase in plant litter production has yet to overcome the influence of the pre‐existing pool of soil organic matter on soil microbial activity and rates of N cycling. Ozone reduces plant litter inputs and also appears to affect the composition of plant litter in a way that reduces microbial biomass and activity.     

Total soil C and N sequestration in a grassland following 10 years of free air CO2 enrichment

Soil C sequestration may mitigate rising levels of atmospheric CO_2. However, it has yet to be determined whether net soil C sequestration occurs in N‐rich grasslands exposed to long‐term elevated CO₂. This study examined whether N‐fertilized grasslands exposed to elevated CO₂ sequestered additional C. For 10 years, Lolium perenne, Trifolium repens, and the mixture of L. perenne/T. repens grasslands were exposed to ambient and elevated CO₂ concentrations (35 and 60 Pa pCO₂). The applied CO₂ was depleted in δ¹³C and the grasslands received low (140 kg ha^?1) and high (560 kg ha^?1) rates of ¹⁵N‐labeled fertilizer. Annually collected soil samples from the top 10 cm of the grassland soils allowed us to follow the sequestration of new C in the surface soil layer. For the first time, we were able to collect dual‐labeled soil samples to a depth of 75 cm after 10 years of elevated CO₂ and determine the total amount of new soil C and N sequestered in the whole soil profile. Elevated CO₂, N‐fertilization rate, and species had no significant effect on total soil C. On average 9.4 Mg new C ha^?1 was sequestered, which corresponds to 26.5% of the total C. The mean residence time of the C present in the 0–10 cm soil depth was calculated at 4.6±1.5 and 3.1±1.1 years for L. perenne and T. repens soil, respectively. After 10 years, total soil N and C in the 0–75 cm soil depth was unaffected by CO₂ concentration, N‐fertilization rate and plant species. The total amount of ¹⁵N‐fertilizer sequestered in the 0–75 cm soil depth was also unaffected by CO₂ concentration, but significantly more ¹⁵N was sequestered in the L. perenne compared with the T. repens swards: 620 vs. 452 kg ha^?1 at the high rate and 234 vs. 133 kg ha^?1 at the low rate of N fertilization. Intermediate values of ¹⁵N recovery were found in the mixture. The fertilizer derived N amounted to 2.8% of total N for the low rate and increased to 8.6% for the high rate of N application. On average, 13.9% of the applied ¹⁵N‐fertilizer was recovered in the 0–75 cm soil depth in soil organic matter in the L. perenne sward, whereas 8.8% was recovered under the T. repens swards, indicating that the N₂‐fixing T. repens system was less effective in sequestering applied N than the non‐N₂‐fixing L. perenne system. Prolonged elevated CO₂ did not lead to an increase in whole soil profile C and N in these fertilized pastures. The potential use of fertilized and regular cut pastures as a net soil C sink under long‐term elevated CO₂ appears to be limited and will likely not significantly contribute to the mitigation of anthropogenic C emissions.