Carbon-based Secondary Compounds at Elevated CO2

From literature sources we compiled the data on carbon-based-secondary compounds CBSC (phenolics and terpenoids) and biomass of 17 plant species grown at different CO₂ concentrations under low and high nutrient availabilities. With a low nutrient availability a possible inverse correlation was found between the biomass and CBSC changes. On the contrary, under a high nutrient availability, both the CBSC and biomass increased with elevated CO₂. The wide variation in the CBSC production among species and compounds (larger responses in phenolics than in terpenoids) indicates that the allocation to CBSC may not completely be governed by changes in CO₂ and nutrient availabilities per se. Yet the comparison shows that elevated CO₂ generally loads the carbon into CBSC [their leaf concentration increased an overall average of 14 % at 700 umol(CO₂) mol-r] which may improve our understanding of the carbon storage and cycling in ecosystems under the “global change” of climate. This revised version was published online in August 2006 with corrections to the Cover Date.     

Elevated CO2 increases belowground respiration in California grasslands

This study was designed to identify potential effects of elevated CO₂ on belowground respiration (the sum of root and heterotrophic respiration) in field and microcosm ecosystems and on the annual carbon budget. We made three sets of respiration measurements in two CO₂ treatments, i.e., (1) monthly in the sandstone grassland and in microcosms from November 1993 to June 1994; (2) at the annual peak of live biomass (March and April) in the serpentine and sandstone grasslands in 1993 and 1994; and (3) at peak biomass in the microcosms with monocultures of seven species in 1993. To help understand ecosystem carbon cycling, we also made supplementary measurements of belowground respiration monthly in sandstone and serpentine grasslands located within 500 m of the CO₂ experiment site. The seasonal average respiration rate in the sandstone grassland was 2.12 mol m^-2 s^-1 in elevated CO₂, which was 42% higher than the 1.49 mol m^-2 s^-1 measured in ambient CO₂ (P=0.007). Studies of seven individual species in the microcosms indicated that respiration was positively correlated with plant biomass and increased, on average, by 70% with CO₂. Monthly measurements revealed a strong seasonality in belowground respiration, being low (0–0.5 mol CO₂ m^-2 s^-1 in the two grasslands adjacent to the CO₂ site) in the summer dry season and high (2–4 mol CO₂ m^-2 s^-1 in the sandstone grassland and 2–7 mol CO₂ m^-2 s^-1 in the microcosms) during the growing season from the onset of fall rains in November to early spring in April and May. Estimated annual carbon effluxes from the soil were 323 and 440 g C m^-2 year^-1 for the sandstone grasslands in ambient and elevated CO₂. That CO₂-stimulated increase in annual soil carbon efflux is more than twice as big as the increase in aboveground net primary productivity (NPP_a) and approximately 60% of NPP_a in this grassland in the current CO₂ environment. The results of this study suggest that below-ground respiration can dissipate most of the increase in photosynthesis stimulated by elevated CO₂.CIWDPB Publication # 1271     

Elevated CO2 alters belowground exoenzyme activities in tussock tundra

A three-year exposure to a CO2 concentration of 680 mol mol-1 altered the enzymic characteristics of root surfaces, associated ectomycorrhizae, and in soils surrounding roots in a tussock tundra ecosystem of north Alaska, USA. At elevated CO2, phosphatase activity was higher on Eriophorum vaginatum root surfaces, ectomycorrhizal rhizomorphs and mantles associated with Betula nana roots, and in Oe and Oi soil horizons associated with plant roots. Also, endocellulase and exocellulase activities at elevated CO2 were higher in ectomycorrhizal rhizomorphs and lower in Oe and Oi soil horizons associated with roots. These results suggest that arctic plants respond to raised CO2 by increasing activities associated with nutrient acquisition, e.g. higher phosphatase activities on surfaces of roots and ectomycorrhizae, and greater cellulase activity in ectomycorrhizae. Changes in enzyme activities of surrounding soils are consistent with an increase in carbon exudation from plant roots, which would be expected to inhibit cellulase activities and stimulate phosphatase activities of soil microflora. These data were used to modify existing simulation models describing tussock phosphatase activities and litter decay. Model projections suggest that observed increases in phosphatase activities at 680 mol mol-1 CO2 could augment total annual phosphorus release within tussocks by more than 40%, at present levels of root and ectomycorrhizae biomass. This includes a nearly three-fold increase in potential phosphatase activity of E. vaginatum roots, per unit of surface area. Observed reductions in cellulase activities could diminish cellulose turnover by 45% in soils within rooting zones, which could substantially increase mineral nitrogen availability in soils due to lowered microbial immobilization.     

Elevated CO2 affects porewater chemistry in a brackish marsh

As atmospheric CO₂ concentrations continue to rise and impact plant communities, concomitant shifts in belowground microbial processes are likely, but poorly understood. We measured monthly porewater concentrations of sulfate, sulfide, methane (CH₄), dissolved inorganic carbon and dissolved organic carbon over a 5-year period in a brackish marsh. Samples were collected using porewater wells (i.e., sippers) in a Schoenoplectus americanus-dominated (C₃ sedge) community, a Spartina patens-dominated (C₄ grass) community and a mixed (C₃ and C₄) community within the marsh. Plant communities were exposed to ambient and elevated (ambient + 340 ppm) CO₂ levels for 15 years prior to porewater sampling, and the treatments continued over the course of our sampling. Sulfate reduction was stimulated by elevated CO₂ in the C₃-dominated community, but not in the C₄-dominated community. Elevated CO₂ also resulted in higher porewater concentrations of CH₄ and dissolved organic carbon in the C₃-dominated system, though inhibition of CH₄ production by sulfate reduction appears to temper the porewater CH₄ response. These patterns mirror the typical divergent responses of C₃ and C₄ plants to elevated CO₂ seen in this ecosystem. Porewater concentrations of nitrogen (as ammonium) and phosphorus did not decrease despite increased plant biomass in the C₃-dominated community, suggesting nutrients do not strongly limit the sustained vegetation response to elevated CO₂. Overall, our data demonstrate that elevated CO₂ drives changes in porewater chemistry and suggest that increased plant productivity likely stimulates microbial decomposition through increases in dissolved organic carbon availability.     

Temperature Effects on Rice at Elevated CO2 Concentration

The continuing increase in atmospheric carbon dioxide concentration([CO₂]) and projections of possible future increases in globalairtemperatures have stimulated interest in the effects of theseclimate variables on agriculturally important food crops. Thisstudywas conducted to determine the effects of [CO₂] and temperatureon rice (Oryza sativa L., cv. IR–30). Rice plants weregrownseason-long in outdoor, naturally sunlit, controlled-environment,plant growth chambers in temperature regimes ranging from 25/18/21°Cto 37/30/34°C (daytime dry bulb air temperature/night-timedry bulb air temperature/paddy water temperature)and [CO₂] of660 µmol CO₂ mol1 air. An ambient chamber was maintainedat a [CO₂] of 330 µmol mol^–1 and temperature regimesof 28/21/25°C. Carbon dioxide enrichment at 28/21/25°Cincreased both biomass accumulation and tillering and increasedgrain yield by 60%. In the 660 µmol mol^–1 [CO₂]treatment, grain yield decreased from 10.4 to 1.0 Mg ha^–1with increasing temperature from 28/21/25°C to the 37/30/34°Ctemperature treatment. Across this temperature range, the numberof panicles plant^–1 nearly doubled while the number ofseeds panicle^–1 declined sharply. These results indicatethat while future increases in atmospheric [CO₂] are likelyto be beneficial to rice growth and yield, potentially largenegative effects on rice yield are possible if air temperaturesalso rise. Key words: Oryza sativa, CO₂, temperature, growth, yield     

Elevated CO2 Affects Predator-Prey Interactions through Altered Performance

Recent research has shown that exposure to elevated carbon dioxide (CO₂) affects how fishes perceive their environment, affecting behavioral and cognitive processes leading to increased prey mortality. However, it is unclear if increased mortality results from changes in the dynamics of predator-prey interactions or due to prey increasing activity levels. Here we demonstrate that ocean pCO₂ projected to occur by 2100 significantly effects the interactions of a predator-prey pair of common reef fish: the planktivorous damselfish Pomacentrus amboinensis and the piscivorous dottyback Pseudochromis fuscus. Prey exposed to elevated CO₂ (880 µatm) or a present-day control (440 µatm) interacted with similarly exposed predators in a cross-factored design. Predators had the lowest capture success when exposed to elevated CO₂ and interacting with prey exposed to present-day CO₂. Prey exposed to elevated CO₂ had reduced escape distances and longer reaction distances compared to prey exposed to present-day CO₂ conditions, but this was dependent on whether the prey was paired with a CO₂ exposed predator or not. This suggests that the dynamics of predator-prey interactions under future CO₂ environments will depend on the extent to which the interacting species are affected and can adapt to the adverse effects of elevated CO₂.     

开放式空气CO2浓度升高与作物/杂草的竞争关系

CO2浓度升高对植物的光合作用、呼吸作用和水分利用等生理过程产生直接影响,进而影响植物的生长繁殖,CO2浓度升高对于具有C3光合途径的植物较具C4光合途径的植物更为有益,由于许多重要的杂草是C4植物,而许多重要的作用是C3植物,CO2浓度升高对杂草/作物的相互关系将有重要影响,本文就全球CO2浓度升高和气候变化对杂草/作物之间竞争关系影响进行综述,同时针对目前研究现状和可持续农业的需要,提出CO2学浓度升高条件下杂草/作物之间竞争关系及未来农田杂草治理方面理论与实践中有待解决的问题。     

CO2 浓度升高对两种沈阳城市森林树种光合特性的影响

利用开顶式气室, 研究了CO2浓度升高条件下城市森林主要树种油松(Pinus tabulaefomis)和银杏(Ginkgo biloba)主要光合特性的变化。结果表明, 整个生长季, CO2浓度升高(700 mmol.mol-1)条件下2树种叶片的净光合速率、可溶性糖、淀粉和可溶性蛋白含量均接近或高于相应对照(自然CO2浓度)值, 但不同树种增加的幅度不同; 而2树种的叶绿素含量和Chl a/Chl b值对CO2浓度升高反应不一, 表现为CO2浓度升高条件下油松的叶绿素含量较对照值高, Chl a/Chl b值降低, 银杏的叶绿素含量为前期升高, 后期降低, Chl a/Chl b值变化与之正好相反, 说明城市森林组成树种对CO2浓度升高的响应具有复杂性。CO2浓度升高条件下, 两树种均未发生光合适应现象。     

Responses of Rice Cultivars to the Elevated CO2

The effect of CO₂ concentration elevated to 575 – 620 µmol mol^–1 on growth, tillering, grain yield, net photosynthetic rate, dark respiration rate, stomatal conductance, sugar content and protein profile of two rice (Oryza sativa L.) cultivars Pusa Basmati-1 and Pusa-677 at flowering stage was studied using open top chambers. The cultivar Pusa Basmati-1 responded more markedly for most of the growth and physiological parameters compared to Pusa-677. The increase in grain yield in Pusa Basmati-1 attributed largely to increased grain number. The increased net photosynthetic rate and greater accumulation of sugar contributed significantly to the accelerated development of leaves and tillers in both the cultivars. The reduction in the low molecular mass proteins including Rubisco and increase in high molecular mass photosystem 2 proteins was observed in both the cultivars. Additional sugars may possibly help in balancing the profile of photosynthetic proteins and sustain greater growth and productivity in rice cultivars.     

Elevated CO2 and temperature impacts on different components of soil CO2 efflux in Douglas-fir terracosms

Although numerous studies indicate that increasing atmospheric CO₂ or temperature stimulate soil CO₂ efflux, few data are available on the responses of three major components of soil respiration [i.e. rhizosphere respiration (root and root exudates), litter decomposition, and oxidation of soil organic matter] to different CO₂ and temperature conditions. In this study, we applied a dual stable isotope approach to investigate the impact of elevated CO₂ and elevated temperature on these components of soil CO₂ efflux in Douglas-fir terracosms. We measured both soil CO₂ efflux rates and the ¹³C and ¹⁸O isotopic compositions of soil CO₂ efflux in 12 sun-lit and environmentally controlled terracosms with 4-year-old Douglas fir seedlings and reconstructed forest soils under two CO₂ concentrations (ambient and 200 ppmv above ambient) and two air temperature regimes (ambient and 4 °C above ambient). The stable isotope data were used to estimate the relative contributions of different components to the overall soil CO₂ efflux. In most cases, litter decomposition was the dominant component of soil CO₂ efflux in this system, followed by rhizosphere respiration and soil organic matter oxidation. Both elevated atmospheric CO₂ concentration and elevated temperature stimulated rhizosphere respiration and litter decomposition. The oxidation of soil organic matter was stimulated only by increasing temperature. Release of newly fixed carbon as root respiration was the most responsive to elevated CO₂, while soil organic matter decomposition was most responsive to increasing temperature. Although some assumptions associated with this new method need to be further validated, application of this dual-isotope approach can provide new insights into the responses of soil carbon dynamics in forest ecosystems to future climate changes.     

Elevated CO2 affects plant responses to variation in boron availability

Aim

Effects of elevated CO₂ on N relations are well studied, but effects on other nutrients, especially micronutrients, are not. We investigated effects of elevated CO₂ on response to variation in boron (B) availability in three unrelated species: seed geranium (Pelargonium x hortorum), barley (Hordeum vulgare), and water fern (Azolla caroliniana).

Methods

Plants were grown at two levels of CO₂ (370, 700?ppm) and low, medium, and high B. Treatment effects were measured on biomass, net photosynthesis (P_n) and related variables, tissue nutrient concentrations, and B transporter protein BOR1.

Results

In geranium, there were interactive effects (P?<?0.05) of B and CO₂ on leaf, stem, and total plant mass, root:shoot ratio, leaf [B], B uptake rate, root [Zn], and P_n. Elevated CO₂ stimulated growth at 45?μM B, but decreased it at 450?μM B and did not affect it at 4.5?μM B. P_n was stimulated by elevated CO₂ only at 45?μM B and chlorophyll was enhanced only at 450?μM B. Soluble sugars increased with high CO₂ only at 4.5 and 45?μM B. High CO₂ decreased leaf [B] and B uptake rate, especially at 450?μM B. Though CO₂ and B individually affected the concentration of several other nutrients, B x CO₂ interactions were evident only for Zn in roots, wherein [Zn] decreased under elevated CO₂. Interactive effects of B and CO₂ on growth were confirmed in (1) barley grown at 0, 30, or 1,000?μM B, wherein growth at high CO₂ was stimulated more at 30?μM B, and (2) Azolla grown at 0, 10, and 1,000?μM B, wherein growth at high CO₂ was stimulated at 0 and 10?μM B.

Conclusion

Thus, low and high B both may limit growth stimulation under elevated vs. current [CO₂], and B deficiency and toxicity, already common, may increase in the future.     

Nematode Genera in Forest Soil Respond Differentially to Elevated CO2

Previous reports suggest that fungivorous nematodes are the only trophic group in forest soils affected by elevated CO₂. However, there can be ambiguity within trophic groups, and we examined data at a genus level to determine whether the conclusion remains similar. Nematodes were extracted from roots and soil of loblolly pine (Pinus taeda) and sweet gum (Liquidambar styraciflua) forests fumigated with either ambient air or CO₂-enriched air. Root length and nematode biomass were estimated using video image analysis. Most common genera included Acrobeloides, Aphelenchoides, Cephalobus, Ditylenchus, Ecphyadorphora, Filenchus, Plectus, Prismatolaimus, and Tylencholaimus. Maturity Index values and diversity increased with elevated CO₂ in loblolly pine but decreased with elevated CO₂ in sweet gum forests. Elevated CO₂ treatment affected the occurrence of more nematode genera in sweet gum than loblolly pine forests. Numbers were similar but size of Xiphinema decreased in elevated CO₂. Abundance, but not biomass, of Aphelenchoides was reduced by elevated CO₂. Treatment effects were apparent at the genus levels that were masked at the trophic level. For example, bacterivores were unaffected by elevated CO₂, but abundance of Cephalobus was affected by CO₂ treatment in both forests.     

Elevated CO2 alters carbon fluxes in early successional Mediterranean ecosystems

Annual carbon budgets of ecosystems are central to our understanding of the biotic control of atmospheric composition, but they are not available under elevated CO₂ for most vegetation types. Using gas exchange techniques, we assessed carbon fluxes of four early successional Mediterranean model communities, consisting of grasses, legumes and composites. The assemblages were grown on the same monoliths for three consecutive years in greenhouses tracking field conditions except for CO₂ maintained at ambient (370 μmol mol^?1) or elevated (700 μmol mol^?1) concentration. During the third year of study, CO₂ enrichment consistently shifted the annual carbon balance towards lower efflux, with displacements between 4.3 and 26.2 mol m^?2 y^?1 (one assemblage became a net CO₂ sink, another just reached equilibrium, and the remaining two remained as a CO₂ source). At least 50% of the shift under elevated CO₂ originated from a decrease in belowground respiration. This indicates that, during this year, CO₂ enrichment did not predominantly enhance C‐cycling, but on the contrary inhibited root respiration or microbial C‐utilization. Although elevated‐CO₂‐grown systems acted as a net CO₂ sink during a longer period of the year (4–7 months) compared with ambient‐CO₂‐grown systems (3–3.5 months), gross canopy photosynthesis was modified only to a limited extent (between ?5.9 and + 14.8%). Interaction between the carbon and the water cycle was apparently responsible for this weak stimulation. In particular, reduced evapotranspiration under elevated CO₂ coincided with inhibited canopy photosynthesis in early spring, most likely resulting from water saturation of the soil. In addition, only the earliest‐planted assemblages had an increased gross canopy photosynthesis during late autumn and early winter. This suggests that a longer summer drought, by delaying the establishment of such an annual type of vegetation, would reduce the positive impact of elevated CO₂ on productivity. Water regime appears to strongly govern the influence of CO₂ on the carbon fluxes in Mediterranean ecosystems with annual herbaceous vegetation.     

Elevated atmospheric CO2 increases microbial growth rates in soil: results of three CO2 enrichment experiments

Increasing the belowground translocation of assimilated carbon by plants grown under elevated CO₂ can cause a shift in the structure and activity of the microbial community responsible for the turnover of organic matter in soil. We investigated the long‐term effect of elevated CO₂ in the atmosphere on microbial biomass and specific growth rates in root‐free and rhizosphere soil. The experiments were conducted under two free air carbon dioxide enrichment (FACE) systems: in Hohenheim and Braunschweig, as well as in the intensively managed forest mesocosm of the Biosphere 2 Laboratory (B2L) in Oracle, AZ. Specific microbial growth rates (μ) were determined using the substrate‐induced respiration response after glucose and/or yeast extract addition to the soil. For B2L and both FACE systems, up to 58% higher μ were observed under elevated vs. ambient CO₂, depending on site, plant species and N fertilization. The μ‐values increased linearly with atmospheric CO₂ concentration at all three sites. The effect of elevated CO₂ on rhizosphere microorganisms was plant dependent and increased for: Brassica napus=Triticum aestivum<Beta vulgaris<Populus deltoides. N deficiency affected microbial growth rates directly (N limitation) and indirectly (changing the quantity of fine roots). So, 50% decrease in N fertilization caused the overall increase or decrease of microbial growth rates depending on plant species. The μ‐value increase was lower for microorganisms growing on yeast extract then for those growing on glucose, i.e. the effect of elevated CO₂ was smoothed on rich vs. simple substrate. So, the r/K strategies ratio can be better revealed by studying growth on simple (glucose) than on rich substrate mixtures (yeast extract). Our results clearly showed that the functional characteristics of the soil microbial community (i.e. specific growth rates) rather than total microbial biomass amount are sensitive to increased atmospheric CO₂. We conclude that the more abundant available organics released by roots at elevated CO₂ altered the ecological strategy of the soil microbial community specifically a shift to a higher contribution of fast‐growing r‐selected species was observed. These changes in functional structure of the soil microbial community may counterbalance higher C input into the soil under elevated atmospheric CO₂ concentration.     

Photosynthesis and Plant Growth at Elevated Levels of CO2

In this review, we discuss the effects of elevated CO₂ levelson photosynthesis in relation to the whole plant growth in terrestrialhigher C₃ plants. Short-term CO₂ enrichment stimulates the rateof photosynthesis. Plant mass is also enhanced by CO₂ enrichment.However, the effects of long-term CO₂ enrichment on photosynthesisare variable. Generally, the prolonged exposure to CO₂ enrichmentreduces the initial stimulation of photosynthesis in many species,and frequently suppresses photosynthesis. These responses areattributed to secondary responses related to either excess carbohydrateaccumulation or decreased N content rather than direct responsesto CO₂. Accumulation of carbohydrates in leaves may lead tothe repression of photosynthetic gene expression and excessstarch seems to hinder CO₂ diffusion. Therefore, the specieswhich have the sink organs for carbohydrate accumulation donot show the suppression of photosynthesis. The suppressionof photosynthesis by CO₂ enrichment is always associated withdecreases in leaf N and Rubisco contents. These decreases arenot due to dilution of N caused by a relative increase in theplant mass but are the result of a decrease in N allocationto leaves at the level of the whole plant, and the decreasein Rubisco content is not selective. Leaf senescence and plantdevelopment are also accelerated by CO₂ enrichment. However,they are independent of each other in some species. Thus, variousresponses to CO₂ observed at the level of a single leaf resultfrom manifold responses at the level of the whole plant grownunder conditions of CO₂ enrichment. (Received July 8, 1999; Accepted August 12, 1999)