Comparison of gas use efficiency and treatment uniformity in a forest ecosystem exposed to elevated [CO2] using pure and prediluted free-air CO2 enrichment technology

A direct comparison of treatment uniformity and CO₂ use of pure and prediluted free-air CO₂ enrichment (FACE) systems was conducted in a forest ecosystem. A vertical release pure CO₂ fumigation system was superimposed on an existing prediluted CO₂ fumigation system and operated on alternate days. The FACE system using prediluted CO₂ fumigation technology exhibited less temporal and spatial variability than the pure CO₂ fumigation system. The pure CO₂ fumigation system tended to over-fumigate the upwind portions of the plot and used 25% more CO₂ than the prediluted CO₂ fumigation system. The increased CO₂ use by the pure CO₂ system was exacerbated at low wind speeds. It is not clear if this phenomenon will also be observed in plots with smaller diameters and low-stature vegetation.     

Stem respiration of Populus species in the third year of free-air CO2 enrichment

Carbon cycling in ecosystems, and especially in forests, is intensively studied to predict the effects of global climate change, and the role which forests may play in 'changing climate change'. One of the questions is whether the carbon balance of forests will be affected by increasing atmospheric CO₂ concentrations. Regarding this question, effects of elevated [CO₂] on woody-tissue respiration have frequently been neglected. Stem respiration of three Populus species ( P. alba L. (Clone 2AS-11), P. nigra L. (Clone Jean Pourtet), and P.  ×  euramericana (Clone I-214)) was measured in a managed, high-density forest plantation exposed to free-air CO₂ enrichment (POPFACE). During the period of measurements, in May of the third year, stem respiration rates were not affected by the FACE treatment. Moreover, FACE did not influence the relationships between respiration rate and both stem temperature and relative growth rate. The results were supported by the reported absence of a FACE-effect on growth and stem wood density.     

Elevated atmospheric CO2 affects soil microbial diversity associated with trembling aspen

The effects of elevated atmospheric CO₂ (560 p.p.m.) and subsequent plant responses on the soil microbial community composition associated with trembling aspen was assessed through the classification of 6996 complete ribosomal DNA sequences amplified from the Rhinelander WI free-air CO₂ and O₃ enrichment (FACE) experiments microbial community metagenome. This in-depth comparative analysis provides an unprecedented, detailed and deep branching profile of population changes incurred as a response to this environmental perturbation. Total bacterial and eukaryotic abundance does not change; however, an increase in heterotrophic decomposers and ectomycorrhizal fungi is observed. Nitrate reducers of the domain bacteria and archaea, of the phylum Crenarchaea , potentially implicated in ammonium oxidation, significantly decreased with elevated CO₂. These changes in soil biota are evidence for altered interactions between trembling aspen and the microorganisms in its surrounding soil, and support the theory that greater plant detritus production under elevated CO₂ significantly alters soil microbial community composition.     

Growth in elevated CO2 enhances temperature response of photosynthesis in wheat

The temperature dependence of C₃ photosynthesis may be altered by the growth environment. The effects of long-term growth in elevated CO₂ on photosynthesis temperature response have been investigated in wheat ( Triticum aestivum L.) grown in controlled chambers with 370 or 700 μmol mol⁻¹ CO₂ from sowing through to anthesis. Gas exchange was measured in flag leaves at ear emergence, and the parameters of a biochemical photosynthesis model were determined along with their temperature responses. Elevated CO₂ slightly decreased the CO₂ compensation point and increased the rate of respiration in the light and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) V_cmax, although the latter effect was reversed at 15°C. With elevated CO₂, J_max decreased in the 15–25°C temperature range and increased at 30 and 35°C. The temperature response (activation energy) of V_cmax and J_max increased with growth in elevated CO₂. CO₂ enrichment decreased the ribulose 1,5-bisphosphate (RuBP)-limited photosynthesis rates at lower temperatures and increased Rubisco- and RuBP-limited rates at higher temperatures. The results show that the photosynthesis temperature response is enhanced by growth in elevated CO₂. We conclude that if temperature acclimation and factors such as nutrients or water availability do not modify or negate this enhancement, the effects of future increases in air CO₂ on photosynthetic electron transport and Rubisco kinetics may improve the photosynthetic response of wheat to global warming.     

Free Air CO2 Enrichment of potato (Solanum tuberosum, L.): design and performance of the CO2-fumigation system

Free Air CO₂ Enrichment (FACE) systems are used to fumigate unconfined field plots with CO₂. As these installations can treat a sufficiently large area without interfering with natural climatic conditions, they are considered important tools for global change research worldwide. However, there is general consensus that elevated capital costs of existing FACE systems as well as high running costs may prevent their application at the required level of scale. A new and small FACE system that was designed to reduce both capital costs and CO₂ use, is described in this paper. Due to its intermediate size (8 m diameter) between the smaller Mini-FACE systems that were developed in Italy and the larger systems designed by the Brookhaven National Laboratory in the USA, it was named Mid-FACE. The Mid-FACE was at first developed as a prototype and then used to enrich field grown potato crops in a CO₂ concentration gradient experimental design. Technical details of a Mid-FACE prototype and of the operational set-up are presented in this paper together with performance data in terms of temporal and spatial control of CO₂ concentrations within the experimental area.     

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

Next generation of elevated [CO2] experiments with crops: a critical investment for feeding the future world

A rising global population and demand for protein-rich diets are increasing pressure to maximize agricultural productivity. Rising atmospheric [CO₂] is altering global temperature and precipitation patterns, which challenges agricultural productivity. While rising [CO₂] provides a unique opportunity to increase the productivity of C₃ crops, average yield stimulation observed to date is well below potential gains. Thus, there is room for improving productivity. However, only a fraction of available germplasm of crops has been tested for CO₂ responsiveness. Yield is a complex phenotypic trait determined by the interactions of a genotype with the environment. Selection of promising genotypes and characterization of response mechanisms will only be effective if crop improvement and systems biology approaches are closely linked to production environments, that is, on the farm within major growing regions. Free air CO₂ enrichment (FACE) experiments can provide the platform upon which to conduct genetic screening and elucidate the inheritance and mechanisms that underlie genotypic differences in productivity under elevated [CO₂]. We propose a new generation of large-scale, low-cost per unit area FACE experiments to identify the most CO₂-responsive genotypes and provide starting lines for future breeding programmes. This is necessary if we are to realize the potential for yield gains in the future.     

Interspecific differences in the response of arbuscular mycorrhizal fungi to Artemisia tridentata grown under elevated atmospheric CO2

Arbuscular mycorrhizal (AM) fungi form mutualistic symbioses with the root systems of most plant species. These mutualisms regulate nutrient exchange in the plant–soil interface and might influence the way in which plants respond to increasing atmospheric CO₂. In other experiments, mycorrhizal responses to elevated CO₂ have been variable, so in this study we test the hypothesis that different genera of AM fungi differ in their response, and in turn alter the plant's response, to elevated CO₂. Four species from three genera of AM fungi were tested. Artemisia tridentata Nutt. seedlings were inoculated with either Glomus intraradices Schenck & Smith, Glomus etunicatum Becker & Gerdemann, Acaulospora sp. or Scutellospora calospora (Nicol. & Gerd.) Walker & Sanders and grown at either ambient CO₂ (350 ppm) or elevated CO₂ (700 ppm). Several significant inter-specific responses were detected. Elevated CO₂ caused percent arbuscular and hyphal colonization to increase for the two Glomus species, but not for Acaulospora sp. or S. calospora . Vesicular colonization was not affected by elevated CO₂ for any fungal species. In the extra-radical phase, the two Glomus species produced a significantly higher number of spores in response to elevated CO₂, whereas Acaulospora sp. and S. calospora developed significantly higher hyphal lengths. These data show that AM fungal taxa differ in their growth allocation strategies and in their responses to elevated CO₂, and that mycorrhizal diversity should not be overlooked in global change research.     

Intraspecific variation in the response of rice (Oryza sativa) to increased CO2– photosynthetic, biomass and reproductive characteristics

Two rice ( Oryza sativa L.) cultivars of contrasting morphologies, IR-36 and Fujiyama-5, were exposed to ambient (360 μl l⁻¹) and ambient plus 300 μl l⁻¹ CO₂ from time of emergence until ca 50% grain fill at the Duke University Phytotron, Durham, North Carolina. Exposure to increased CO₂ resulted in about a 50% increase in the photosynthetic rate for both cultivars and photosynthetic enhancement was still evident after 3 months of exposure to a high CO₂ environment. The photosynthetic response at 5% CO₂ and the response of CO₂ assimilation (A) to internal CO₂ (C_i) suggest a reallocation of biochemical resources from RuBP carboxylation to RuBP regeneration. Increases in total plant biomass at elevated CO₂ were approximately the same in both cultivars, although differences in allocation patterns were noted in root/shoot ratio. Differences in reproductive characteristics were also observed between cultivars at an elevated CO₂ environment with a significant increase in harvest index for IR-36 but not for Fujiyama-5. Changes in carbon allocation in reproduction between these two cultivars suggest that lines of rice could be identified that would maximize reproductive output in a future high CO₂ environment.     

Wood composition and energy content in a poplar short rotation plantation on fertilized agricultural land in a future CO2 atmosphere

To study the influence of elevated CO₂ and nitrogen (N) fertilization on wood properties and energy, Populus × euramericana trees were exposed to ambient CO₂ (about 370 μmol mol⁻¹ CO₂) or elevated CO₂ (about 550 μmol mol⁻¹ CO₂) using Free Air CO₂ Enrichment (FACE) technology in combination with two N levels. Elevated CO₂ was maintained for 5 years. After three growing seasons, the plantation was coppiced, one half of each experimental plot was fertilized and secondary sprouts were harvested after two growing seasons. Fourier transform infrared (FT-IR) spectra of wood revealed significant effects of both elevated CO₂ and N fertilization on wood chemistry, in particular, significant increases in lignin and decreases in N content. These results were corroborated by chemical analysis. Neither elevated CO₂ nor N fertilization affected the calorific value of wood, which was 19.3 MJ kg⁻¹. N fertilization enhanced the energy production per land area by 16–69% because of higher aboveground woody biomass production than on nonfertilized land. Estimates indicate that high yielding poplar short rotation cultivation may significantly contribute as an alternative feedstock for energy production.     

Productivity and community structure of ectomycorrhizal fungal sporocarps under increased atmospheric CO2 and O3

Sporocarp production is essential for ectomycorrhizal fungal recombination and dispersal, which influences fungal community dynamics. Increasing atmospheric carbon dioxide (CO₂) and ozone (O₃) affect host plant carbon gain and allocation, which may in turn influence ectomycorrhizal sporocarp production if the carbon available to the ectomycorrhizal fungus is dependant upon the quantity of carbon assimilated by the host. We measured sporocarp production of ectomycorrhizal fungi over 4 years at the Aspen FACE (free air CO₂ enrichment) site, which corresponded to stand ages seven to 10 years. Total mean sporocarp biomass was greatest under elevated CO₂, regardless of O₃ concentration, while it was generally lowest under elevated O₃ with ambient CO₂. Community composition differed significantly among the treatments, with less difference in the final year of the study. Whether this convergence was due to succession or environmental factors is uncertain. CO₂ and O₃ affect ectomycorrhizal sporocarp productivity and community composition, with likely effects on dispersal, colonization and sporocarp-dependent food webs.     

Mitigation of adverse effects of rising CO2 on a planktonic herbivore by mixed algal diets

Putative future increase in atmospheric CO₂ is expected to adversely affect herbivore growth due to decrease in contents of key nutrients such as nitrogen and phosphorus (P) relative to carbon in primary producers including plant and algal species. However, as many herbivores are polyphagous and as the response of primary producers to elevated CO₂ is highly species-specific, effects of elevated CO₂ on herbivore growth may differ between feeding conditions with monospecific and multiproducer diets. To examine this possibility, we performed CO₂ manipulation experiments under a P-limited condition with a planktonic herbivore, Daphnia , and three algal species, Scenedesmus obliquus (green algae), Cyclotella sp. (diatoms) and Synechococcus sp. (cyanobacteria). Semibatch cultures with single algal species (monocultures) and multiple algal species (mixed cultures) were grown at ambient (360 ppm) and high CO₂ levels (2000 ppm) that were within the natural range in lakes. Both in the mono- and mixed cultures, algal steady state abundance increased but algal P : C and N : C ratios decreased when they were grown at high CO₂. As expected, Daphnia fed monospecific algae cultured at high CO₂ had decreased growth rates despite increased algal abundance. However, when fed mixed algae cultured at high CO₂, especially consisting of diatoms and cyanobacteria or the three algal species, Daphnia maintained high growth rates despite lowered P and N contents relative to C in the algal diets. These results imply that algal diets composed of multiple species can mitigate the adverse effects of elevated CO₂ on herbivore performance, although the magnitude of this mitigation depends on the composition of algal species involved in the diets.     

The interaction of high temperature and elevated CO2 on photosynthetic acclimation of single leaves of rice in situ

Rice ( Oryza sativa L. cv. IR72) was grown at three different CO₂ concentrations (ambient, ambient + 200 μmol mol⁻¹, ambient + 300 μmol mol⁻¹) at two different growth temperatures (ambient, ambient + 4°C) from sowing to maturity to determine longterm photosynthetic acclimation to elevated CO₂ with and without increasing temperature. Single leaves of rice showed a cooperative enhancement of photosynthetic rate with elevated CO₂ and temperature during tillering, relative to the elevated CO₂ condition alone. However, after flowering, the degree of photosynthetic stimulation by elevated CO₂ was reduced for the ambient + 4°C treatment. This increasing insensitivity to CO₂ appeared to be accompanied by a reduction in ribulose-1.5-bisphosphate carboxylase/oxygenase (Rubisco) activity and/or concentration as evidenced by the reduction in the assimilation (A) to internal CO₂ (C₁) response curve. The reproductive response (e.g. percent filled grains, panicle weight) was reduced at the higher growth temperature and presumably reflects a greater increase in floral sterility. Results indicate that while CO₂ and temperature could act synergistically at the biochemical level, the direct effect of temperature on floral development with a subsequent reduction in carbon utilization may change sink strength so as to limit photosynthetic stimulation by elevated CO₂ concentration.     

Contrasting effects of climate and CO2 on Amazonian ecosystems since the last glacial maximum

The nature of Amazonian ecosystem responses to the large-scale environmental changes characterizing glacial–interglacial cycles is poorly understood. We investigated this issue with a series of transient, continuous 21 000-year simulations using a dynamic process-based ecosystem model. Our results indicate that the Amazon Basin has been dominated by evergreen rain forests since the last glacial maximum (LGM), demonstrating the resilience of this ecosystem to glacial–interglacial environmental change. We find that biome shifts in ecotonal areas since the LGM were driven predominantly by climate change, while coincident, increased ecosystem carbon storage throughout the Amazon Basin was driven largely by CO₂. Our findings imply that recent observed biomass increases in contemporary rain forest plots might be part of a long-term trend driven by the anthropogenic rise in CO₂ over recent centuries.     

Rice production in a changing climate: a meta-analysis of responses to elevated carbon dioxide and elevated ozone concentration

Rice is arguably the most important food source on the planet and is consumed by over half of the world's population. Considerable increases in yield are required over this century to continue feeding the world's growing population. This meta-analysis synthesizes the research to date on rice responses to two elements of global change, rising atmospheric carbon dioxide concentration ([CO₂]) and rising tropospheric ozone concentration ([O₃]). On an average, elevated [CO₂] (627 ppm) increased rice yields by 23%. Modest increases in grain mass and larger increases in panicle and grain number contributed to this response. The response of rice to elevated [CO₂] varied with fumigation technique. The more closely the fumigation conditions mimicked field conditions, the smaller was the stimulation of yield by elevated [CO₂]. Free air concentration enrichment (FACE) experiments showed only a 12% increase in rice yield. The rise in atmospheric [CO₂] will be accompanied by increases in tropospheric O₃ and temperature. When compared with rice grown in charcoal-filtered air, rice exposed to 62 ppb O₃ showed a 14% decrease in yield. Many determinants of yield, including photosynthesis, biomass, leaf area index, grain number and grain mass, were reduced by elevated [O₃]. While there have been too few studies of the interaction of CO₂ and O₃ for meta-analysis, the interaction of temperature and CO₂ has been studied more widely. Elevated temperature treatments negated any enhancement in rice yield at elevated [CO₂], which suggests that identifying high temperature tolerant germplasm will be key to realizing yield benefits in the future.     

The role of temperature in determining the stimulation of CO2 assimilation at elevated carbon dioxide concentration in soybean seedlings

Soybean ( Glycine max cv. Clark) was grown at both ambient (ca 350 μmol mol⁻¹) and elevated (ca 700 μmol mol⁻¹) CO₂ concentration at 5 growth temperatures (constant day/night temperatures of 20, 25, 30, 35 and 40°C) for 17–22 days after sowing to determine the interaction between temperature and CO₂ concentration on photosynthesis (measured as A, the rate of CO₂ assimilation per unit leaf area) at both the single leaf and whole plant level. Single leaves of soybean demonstrated increasingly greater stimulation of A at elevated CO₂ as temperature increased from 25 to 35°C (i.e. optimal growth rates). At 40°C, primary leaves failed to develop and plants eventually died. In contrast, for both whole plant A and total biomass production, increasing temperature resulted in less stimulation by elevated CO₂ concentration. For whole plants, increased CO₂ stimulated leaf area more as growth temperature increased. Differences between the response of A to elevated CO₂ for single leaves and whole plants may be related to increased self-shading experienced by whole plants at elevated CO₂ as temperature increased. Results from the present study suggest that self-shading could limit the response of CO₂ assimilation rate and the growth response of soybean plants if temperature and CO₂ increase concurrently, and illustrate that light may be an important consideration in predicting the relative stimulation of photosynthesis by elevated CO₂ at the whole plant level.     

Species-specific responses to atmospheric carbon dioxide and tropospheric ozone mediate changes in soil carbon

We repeatedly sampled the surface mineral soil (0–20 cm depth) in three northern temperate forest communities over an 11-year experimental fumigation to understand the effects of elevated carbon dioxide (CO₂) and/or elevated phyto-toxic ozone (O₃) on soil carbon (C). After 11 years, there was no significant main effect of CO₂ or O₃ on soil C. However, within the community containing only aspen ( Populus tremuloides Michx.), elevated CO₂ caused a significant decrease in soil C content. Together with the observations of increased litter inputs, this result strongly suggests accelerated decomposition under elevated CO_2. In addition, an initial reduction in the formation of new (fumigation-derived) soil C by O₃ under elevated CO₂ proved to be only a temporary effect, mirroring trends in fine root biomass. Our results contradict predictions of increased soil C under elevated CO₂ and decreased soil C under elevated O₃ and should be considered in models simulating the effects of Earth's altered atmosphere.     

Elevated CO2 enhances stomatal responses to osmotic stress and abscisic acid in Arabidopsis thaliana

A , carbon assimilation rate
ABA, abscisic acid
Ci , intercellular space CO₂ concentration
g , leaf conductance
WUE, water use efficiency

Carbon dioxide and abscisic acid (ABA) are two major signals triggering stomatal closure. Their putative interaction in stomatal regulation was investigated in well-watered air-grown or double CO₂-grown Arabidopsis thaliana plants, using gas exchange and epidermal strip experiments. With plants grown in normal air, a doubling of the CO₂ concentration resulted in a rapid and transient drop in leaf conductance followed by recovery to the pre-treatment level after about two photoperiods. Despite the fact that plants placed in air or in double CO₂ for 2 d exhibited similar levels of leaf conductance, their stomatal responses to an osmotic stress (0·16–0·24 MPa) were different. The decrease in leaf conductance in response to the osmotic stress was strongly enhanced at elevated CO₂. Similarly, the drop in leaf conductance triggered by 1 μ M ABA applied at the root level was stronger at double CO₂. Identical experiments were performed with plants fully grown at double CO₂. Levels of leaf conductance and carbon assimilation rate measured at double CO₂ were similar for air-grown and elevated CO₂-grown plants. An enhanced response to ABA was still observed at high CO₂ in pre-conditioned plants. It is concluded that: (i) in the absence of stress, elevated CO₂ slightly affects leaf conductance in A. thaliana ; (ii) there is a strong interaction in stomatal responses to CO₂ and ABA which is not modified by growth at elevated CO₂.     

Vegetation Restoration and Its Effects on Carbon Balance in Guangdong Province, China

The significance of the regional carbon (C) balance of vegetation restoration to global change was studied in Guangdong Province, one of the most populated provinces in China. The percentage of the province in forestland cover increased steadily from 26.23% in 1979 to 50.11% in 1998 owing to restoration of forests. During this period, the area increase in the conifer forest was 424.83 × 10⁴ ha, whereas the area in broad-leaved, mixed, and other forests increased by 68.82 × 10⁴, 18.94 × 10⁴, and 60.46 × 10⁴ ha, respectively. C sequestration by forest soils from 1979 to 1998 was 389.36 Mt, equivalent to 1.43 Pg fixed carbon dioxide (CO₂). The total annual CO₂ sequestration by forests and the soils was 118.05 Mt, which was about half of the annual CO₂ emission in Guangdong. The role of forest management and restoration in improving the carbon balance is discussed.     

A field fumigation system for elevated carbon dioxide exposure in chaparral shrubs

1. Modifications and improvements in the hardware and software of a free-air CO₂ enrichment (FACE) system are described. The modifications were undertaken to minimize the amount of structure required and to improve software control of CO₂ set points.
2. A new low-cost CO₂ flow controller which provides increased reliability is described.
3. Analysis of system performance during a 79-day evaluation period of operation (13 h day^–1) showed that for 1 min averages, the system is capable of maintaining a 550 p.p.m. ± 10% CO₂ set point during 78% of the operating time and a 550 p.p.m. ± 20% set point during 95% of the operating time. Ten-minute averages were within ± 10% and 20% during 87% and 96% of the operating time, respectively.
4. Continuous measurements of the spatial variation in CO₂ concentration inside the FACE ring over an 18-day period showed that of the total 16-m diameter treatment area, the central 11-m diameter portion remains within the 550 p.p.m. ± 10% set point.
5. Daily course gas-exchange measurements in matched individuals of the chaparral shrub Adenostoma fasciculatum just prior to and following a 6-week fumigation period at 550 p.p.m. CO₂ in the FACE treatment ring showed the FACE plants with reduced photosynthetic rates and higher (less stressful) water potentials compared with control A. fasciculatum plants measured at the same times and conditions, indicating the responsiveness of this species to elevated CO₂ conditions.