Production, turnover and mycorrhizal colonization of root systems of three Populus species grown under elevated CO2 (POPFACE)

A fast growing high density Populus plantation located in central Italy was exposed to elevated carbon dioxide for a period of three years. An elevated CO₂ treatment (550 ppm), of 200 ppm over ambient (350 ppm) was provided using a FACE technique. Standing root biomass, fine root turnover and mycorrhizal colonization of the following Populus species was examined: Populus alba L., Populus nigra L., Populus x euramericana Dode (Guinier). Elevated CO₂ increased belowground allocation of biomass in all three species examined, standing root biomass increased by 47–76% as a result of FACE treatment. Similarly, fine root biomass present in the soil increased by 35–84%. The FACE treatment resulted in 55% faster fine root turnover in P. alba and a 27% increase in turnover of roots of P. nigra and P. x euramericana. P. alba and P. nigra invested more root biomass into deeper soil horizon under elevated CO₂. Response of the mycorrhizal community to elevated CO₂ was more varied, the rate of infection increased only in P. alba for both ectomycorrhizal (EM) and arbuscular mycorrhizas (AM). The roots of P. nigra showed greater infection only by AM and the colonization of the root system of P. x euramericana was not affected by FACE treatment. The results suggest that elevated atmospheric CO₂ conditions induce greater belowground biomass investment, which could lead to accumulation of assimilated C in the soil profile. This may have implications for C sequestration and must be taken into account when considering long‐term C storage in the soil.     

Stomatal conductance and not stomatal density determines the long-term reduction in leaf transpiration of poplar in elevated CO<Subscript>2</Subscript>

Using a free-air CO₂ enrichment (FACE) experiment, poplar trees (Populus × euramericana clone I214) were exposed to either ambient or elevated [CO₂] from planting, for a 5-year period during canopy development, closure, coppice and re-growth. In each year, measurements were taken of stomatal density (SD, number mm⁻²) and stomatal index (SI, the proportion of epidermal cells forming stomata). In year 5, measurements were also taken of leaf stomatal conductance (g _s, μmol m⁻² s⁻¹), photosynthetic CO₂ fixation (A, mmol m⁻² s⁻¹), instantaneous water-use efficiency (A/E) and the ratio of intercellular to atmospheric CO₂ (C_i:C_a). Elevated [CO₂] caused reductions in SI in the first year, and in SD in the first 2 years, when the canopy was largely open. In following years, when the canopy had closed, elevated [CO₂] had no detectable effects on stomatal numbers or index. In contrast, even after 5 years of exposure to elevated [CO₂], g _s was reduced, A/E was stimulated, and C_i:C_a was reduced relative to ambient [CO₂]. These outcomes from the long-term realistic field conditions of this forest FACE experiment suggest that stomatal numbers (SD and SI) had no role in determining the improved instantaneous leaf-level efficiency of water use under elevated [CO₂]. We propose that altered cuticular development during canopy closure may partially explain the changing response of stomata to elevated [CO₂], although the mechanism for this remains obscure.     

Carbon-based secondary metabolites and internal nitrogen pools in Populus nigra under Free Air CO2 Enrichment (FACE) and nitrogen fertilisation

The goal of this study was to investigate whether increased nitrogen use efficiency found in Populus nigra L. (Jean Pourtet) under elevated CO₂ would correlate with changes in the production of carbon-based secondary compounds (CBSCs). Using Free-Air CO₂ Enrichment (FACE) technology, a poplar plantation was exposed to either ambient (about 370 μmol mol⁻¹ CO₂) or elevated (about 550 μmol mol⁻¹ CO₂) [CO₂] for 5 years. After three growing seasons, the plantation was coppiced and half of the experimental plots were fertilized with nitrogen. CBSCs, total nitrogen and lignin-bound nitrogen were quantified in secondary sprouts in seasons of active growth and dormancy during 2 years after coppicing. Neither elevated CO₂ nor nitrogen fertilisation alone or in combination influenced lignin concentrations in wood. Soluble phenolics and soluble proteins in wood decreased slightly in response to elevated CO₂. Higher nitrogen supply stimulated formation of CBSCs and increased protein concentrations. Lignin-bound nitrogen in wood ranged from 0.37–1.01 mg N g⁻¹ dry mass accounting for 17–26% of total nitrogen in wood, thus, forming a sizeable nitrogen fraction resistant to chemical degradation. The concentration of this nitrogen fraction was significantly decreased by elevated CO₂, increased in response to nitrogen fertilisation and showed a significant CO₂ × fertilisation interaction. Seasonal changes markedly affected the internal nitrogen pools. Soluble proteins in wood were 52–143% higher in the dormant than in the growth season. Positive correlations existed between the biosynthesis of proteins and CBSCs. The limited responses to elevated CO₂ and nitrogen fertilisation indicate that growth and defence are well orchestrated in P. nigra and that changes in the balance of both resources—nitrogen and C—have only marginal effects on wood chemistry. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Leaf area is stimulated in Populus by free air CO2 enrichment (POPFACE), through increased cell expansion and production

The effects of free‐air CO₂ enrichment (FACE) on leaf growth in Populus, was studied. For the first time in field conditions, both the production and expansion of leaf cells were shown to be sensitive to atmospheric carbon dioxide. Leaf area expansion rate and final leaf size were stimulated under FACE for three species (Populus x euramericana (I‐214), P. nigra (Jean Pourtet) and P. alba (2AS‐11), with the largest effect observed for P. x euramericana (61%). In this species and in P. nigra, both epidermal cell size and cell number were increased, whereas for P. alba, only cell production was increased in FACE. Two findings suggest that changes in the cell wall may be important in explaining larger leaf cells in FACE: (i) Leaf cell wall extensibility of rapidly growing leaves increased in all species in FACE; and (ii) an increase in xyloglucan endotransglycosylase activity, a cell wall‐loosening enzyme, was increased in FACE and associated with leaf growth rate. The results suggest that the mechanisms by which FACE promotes leaf growth differ, depending on species. Despite this, increases in final leaf size provide an important component driving increased biomass accumulation in POPFACE, during this first year of rapid growth, prior to canopy closure. The question as to whether these effects are the result of a direct response to CO₂, or are driven indirectly through substrate availability remains unresolved, although evidence from the literature suggests that the latter mechanism is most likely.     

Gross primary production is stimulated for three Populus species grown under free-air CO2 enrichment from planting through canopy closure

How forests will respond to rising [CO₂] in the long term is uncertain, most studies having involved juvenile trees in chambers prior to canopy closure. Poplar free‐air CO₂ enrichment (Viterbo, Italy) is one of the first experiments to grow a forest from planting through canopy closure to coppice, entirely under open‐air conditions using free‐air CO₂ enrichment technology. Three Populus species: P. alba, P. nigra and P. x euramericana, were grown in three blocks, each containing one control and one treatment plot in which CO₂ was elevated to the expected 2050 concentration of 550 ppm. The objective of this study was to estimate gross primary production (GPP) from recorded leaf photosynthetic properties, leaf area index (LAI) and meteorological conditions over the complete 3‐year rotation cycle. From the meteorological conditions recorded at 30 min intervals and biweekly measurements of LAI, the microclimate of leaves within the plots was estimated with a radiation transfer and energy balance model. This information was in turn used as input into a canopy microclimate model to determine light and temperature of different leaf classes at 30 min intervals which in turn was used with the steady‐state biochemical model of leaf photosynthesis to compute CO₂ uptake by the different leaf classes. The parameters of these models were derived from measurements made at regular intervals throughout the coppice cycle. The photosynthetic rates for different leaf classes were summed to obtain canopy photosynthesis, i.e. GPP. The model was run for each species in each plot, so that differences in GPP between species and treatments could be tested statistically. Significant stimulation of GPP driven by elevated [CO₂] occurred in all 3 years, and was greatest in the first year (223–251%), but markedly lower in the second (19–24%) and third years (5–19%). Increase in GPP in elevated relative to control plots was highest for P. nigra in 1999 and for P. x euramericana in 2000 and 2001, although in 1999 P. alba had a higher GPP than P. x euramericana. Our analysis attributed the decline in stimulation to canopy closure and not photosynthetic acclimation. Over the 3‐year rotation cycle from planting to harvest, the cumulative GPP was 4500, 4960 and 4010 g C m^?2 for P. alba, P. nigra and P. x euramericana, respectively, in current [CO₂] and 5260, 5800 and 5000 g C m^?2 in the elevated [CO₂] treatments. The relative changes were consistent with independent measurements of net primary production, determined independently from biomass increments and turnover.     

Does elevated atmospheric CO2 concentrations affect wood decomposition?

This study was conducted to test the hypothesis that wood tissues generated under elevated atmospheric [CO₂] have lower quality and subsequent reduced decomposition rates. Chemical composition and subsequent field decomposition rates were studied for beech (Fagus sylvatica L.) twigs grown under ambient and elevated [CO₂] in open top chambers. Elevated [CO₂] significantly affected the chemical composition of beech twigs, which had 38% lower N and 12% lower lignin concentrations than twigs grown under ambient [CO₂]. The strong decrease in N concentration resulted in a significant increase in the C/N and lignin/N ratios of the beech wood grown at elevated [CO₂]. However, the elevated [CO₂] treatment did not reduce the decomposition rates of twigs, neither were the dynamics of N and lignin in the decomposing beech wood affected by the [CO₂] treatment, despite initial changes in N and lignin concentrations between the ambient and elevated [CO₂] beech wood. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of elevated CO<Subscript>2</Subscript> and nitrogen on wood structure related to water transport in seedlings of two deciduous broad-leaved tree species

Wood structure might be altered through the physiological responses to atmospheric carbon dioxide concentration ([CO₂]) and nitrogen (N) deposition. We investigated growth, water relations and wood structure of 1-year-old seedlings of two deciduous broad-leaved tree species, Quercus mongolica (oak, a ring-porous species) and Alnus hirsuta (alder, a diffuse-porous species and N₂–fixer), grown under a factorial combination of two levels of [CO₂] (36 and 72 Pa) and nitrogen supply (N; low and high) for 141 days in phytotron chambers. In oak, there was no significant effect of [CO₂] on wood structure, although elevated [CO₂] tended to decrease stomatal conductance (g _s) and increased water use efficiency regardless of the N treatment. However, high N supply increased root biomass and induced wider earlywood and larger vessels in the secondary xylem in stems, leading to increased hydraulic conductance. In alder, there was significant interactive effect of [CO₂] and N on vessel density, and high N supply increased the mean vessel area. Our results suggest that wood structures related to water transport were not markedly altered, although elevated [CO₂] induced changes in physiological parameters such as g _s and biomass allocation, and that N fertilization had more pronounced effects on non-N₂-fixing oak than on N₂-fixing alder.     

Woody biomass production during the second rotation of a bio-energy Populus plantation increases in a future high CO2 world

The quickly rising atmospheric carbon dioxide (CO₂)‐levels, justify the need to explore all carbon (C) sequestration possibilities that might mitigate the current CO₂ increase. Here, we report the likely impact of future increases in atmospheric CO₂ on woody biomass production of three poplar species (Populus alba L. clone 2AS‐11, Populus nigra L. clone Jean Pourtet and Populus×euramericana clone I‐214). Trees were growing in a high‐density coppice plantation during the second rotation (i.e., regrowth after coppice; 2002–2004; POPFACE/EUROFACE). Six plots were studied, half of which were continuously fumigated with CO₂ (FACE; free air carbon dioxide enrichment of 550 ppm). Half of each plot was fertilized to study the interaction between CO₂ and nutrient fertilization. At the end of the second rotation, selective above‐ and belowground harvests were performed to estimate the productivity of this bio‐energy plantation. Fertilization did not affect growth of the poplar trees, which was likely because of the high rates of fertilization during the previous agricultural land use. In contrast, elevated CO₂ enhanced biomass production by up to 29%, and this stimulation did not differ between above‐ and belowground parts. The increased initial stump size resulting from elevated CO₂ during the first rotation (1999–2001) could not solely explain the observed final biomass increase. The larger leaf area index after canopy closure and the absence of any major photosynthetic acclimation after 6 years of fumigation caused the sustained CO₂‐induced biomass increase after coppice. These results suggest that, under future CO₂ concentrations, managed poplar coppice systems may exhibit higher potential for C sequestration and, thus, help mitigate climate change when used as a source of C‐neutral energy.     

Three years of free-air CO2 enrichment (POPFACE) only slightly affect profiles of light and leaf characteristics in closed canopies of Populus

Modelling is used to predict long‐term forest responses to increased atmospheric CO₂ concentrations. Although productivity models are based on light intercepted by the canopy, very little experimental data are available for closed forest stands. Nevertheless, the relationships between light inside a canopy, leaf area, canopy structure, and individual leaf characteristics may be affected by elevated CO₂, affecting in turn carbon gain. Using a free‐air CO₂ enrichment (FACE) design in a high‐density plantation of Populus spp., we studied the effects of increased CO₂ concentrations on transmittance (τ) of photosynthetic photon flux density (Q_p), on ratios of red/far‐red light (R/FR), on leaf area index (LAI), on leaf inclination, on leaf chlorophyll (chl) and nitrogen (N) concentrations, and on specific leaf area (SLA) in the 2nd and 3rd years of treatment. Continuous measurements of τ were made in addition to canopy height profiles of light and leaf characteristics. Two years of Q_p measurements showed an average decrease of canopy transmittance in the FACE treatment, with very small differences at canopy closure. Results were explained by an unaffected LAI in closed canopies, without a FACE‐induced stimulation of relative crown depth. In agreement, leaf inclination and extinction coefficients for light were similar in control and FACE conditions. Ratios of R/FR were not significantly affected by the FACE treatment, neither were leaf characteristics, with the exception of leaf N, which allows speculation about N limitation. In general, treatment differences in canopy profiles resulted from an initial stimulation of height growth in the FACE treatment. P. × euramericana differed from P. alba and P. nigra, but species did not differ significantly in their response to the FACE treatment. By the time fast‐growing high‐density forest plantations have passed the exponential growth phase and reached canopy closure, the likely effects of elevated atmospheric CO₂ concentration on canopy architecture and absorption of Q_p are minor.     

Ribulose-1,5-bisphosphate regeneration limitation in rice leaf photosynthetic acclimation to elevated CO2

Our previous study has demonstrated that both RuBP carboxylation limitation and RuBP regeneration limitation exist simultaneously in rice grown under free-air CO₂ enrichment (FACE, about 200 μmol mol⁻¹ above the ambient air CO₂ concentration) conditions [G.-Y. Chen, Z.-H. Yong, Y. Liao, D.-Y. Zhang, Y. Chen, H.-B. Zhang, J. Chen, J.-G. Zhu, D.-Q. Xu, Photosynthetic acclimation in rice leaves to free-air CO₂ enrichment related to both ribulose-1,5-bisphosphate carboxylase limitation and ribulose-1,5-bisphosphate regeneration limitation. Plant Cell Physiol. 46 (2005) 1036–1045]. To explore the mechanism for forming of RuBP regeneration limitation, we conducted the gas exchange measurements and some biochemical analyses in FACE-treated and ambient rice plants. Net CO₂ assimilation rate (A_net) in FACE leaves was remarkably lower than that in ambient leaves when measured at the same CO₂ concentration, indicating that photosynthetic acclimation to elevated CO₂ occurred. In the meantime the maximum electron transport rate (ETR) (J_max), maximum carboxylation rate (V_cmax) in vivo, and RuBP contents decreased significantly in FACE leaves. The whole chain electron transport rate and photophosphorylation rate reduced significantly while ETR of photosystem II (PSII) did not significantly decrease and ETR of photosystem I (PSI) was significantly increased in the chloroplasts from FACE leaves. Further, the amount of cytochrome (Cyt) f protein, a key component localized between PSII and PSI, was remarkably declined in FACE leaves. It appears that during photosynthetic acclimation the decline in the Cyt f amount is an important cause for the decreased RuBP regeneration capacity by decreasing the whole chain electron transport in FACE leaves.     

Nitrogen and Carbon Concentrations, and Stable Isotope Ratios in Mediterranean Shrubs Growing in the Proximity of a CO2 spring

Seasonal changes in foliage nitrogen (N) and carbon (C) concentrations and δ¹⁵N and δ¹³C ratios were monitored during a year in Erica arborea, Myrtus communis and Juniperus communis co-occurring at a natural CO₂ spring (elevated [CO₂], about 700 μmol mol⁻¹) and at a nearby control site (ambient [CO₂], 360 μmol mol⁻¹) in a Mediterranean environment. Leaf N concentration was lower in elevated [CO₂] than in ambient [CO₂] for M. communis, higher for J. communis, and dependent on the season for E. arborea. Leaf C concentration was negatively affected by atmospheric CO₂ enrichment, regardless of the species. C/N ratio varied concomitantly to N. Leaves in elevated [CO₂] showed lower δ¹³C, and therefore likely lower water use efficiencies than leaves at the control site, regardless of the species, suggesting substantial photosynthetic acclimation under long-term CO₂-enriched atmosphere. Leaves of E. arborea showed lower values of δ¹⁵N under elevated [CO₂], but this was not the case of M. communis and J. communis foliage. The use of the resources and leaf chemical composition are affected by elevated [CO₂], but such an effect varies during the year, and is species-dependent. The seasonal dependency and species specificity suggest that plants are able to exploit different available water and N resources within Mediterranean sites. This revised version was published online in July 2006 with corrections to the Cover Date.     

Elevated CO2 concentration and temperature effects on the partitioning of chemical components along juvenile Scots pine stems (Pinus sylvestris L.)

The effects of doubled ambient [CO₂] and different temperature levels on young Pinus sylvestris growing in phytotron chambers were studied. Five chambers were supplied with ~380 (‘ambient air’) and five with ~700 μmol mol⁻¹ CO₂ (‘elevated [CO₂]’). Temperature levels in the chambers ranged in increment steps of 2°C from −4°C to +4°C relative to the long-term monthly (day and night) average air temperature levels in Berlin–Dahlem. Substrate was medium fertile; soil moisture and air humidity were kept constant. After three vegetation periods twigs and stems were harvested, weighed, homogenized, and analyzed chemically. There was no significant temperature effect on wood mass accumulation, clearest positive [CO₂] effect occurred in the youngest twigs. In total, wood mass increased by 28.5% at doubled ambient [CO₂]. N-contents (percentage) decreased at elevated [CO₂] in the uppermost stem sections and not in twig wood causing wider C/N ratios in total. In response to elevated temperature, N-contents decreased slightly in twigs (~0.3%). Traces of free glucose, fructose and sucrose, which decreased from the top to the bottom, were found in stem wood, in contrast to traces of starch that increased from the top to the bottom. In response to elevated [CO₂] only a little more (0.05%) was accumulated in the top shoot and in tendency; glucose, fructose, and sucrose contents were lower at the bottom of stems as compared to the control. There was no obvious response of these non-structural carbohydrates to elevated temperature except for starch that decreased to half of the content from the lowest to the highest temperature level. Among the hemicellulose compounds, rhamnose and arabinose declined from the top shoot to the bottom of stem, whereas 4-O-methyl-d-glucuronic-acid, mannose, and xylose increased. Contents (percentage) of galactose remained approximately stable along the stem. The clearest positive effect of elevated [CO₂] along the whole stem was found for mannose with differences of 0.6–0.3%. In contrast to rhamnose and arabinose that showed a negative response to elevated [CO₂], mannose was reduced towards the uppermost stem sections. The 4-O-methyl-d-glucuronic-acid was slightly lowered at the bottom, and galactose and xylose showed no [CO₂] response. The only hemicellulose compound which reacted to temperature elevation was galactose. It increased slightly (~0.1% per 1°C). Cellulose and lignin (Klason) behaved oppositely: cellulose increased and lignin decreased from the top to the bottom. These structural components behaved reversely also in response to elevated [CO₂]. In stem parts above the bottom section, cellulose content was slightly higher at elevated [CO₂], and lignin content was slightly lower at the bottom. Lignin reacted to temperature elevation by a very slight increase on the average (~0.1% per one 1°C). Cellulose, however, decreased by ~0.2% per 1°C temperature elevation. The importance of persistent sinks of carbon in woody plant parts is discussed in respect to the greenhouse effect.     

Leaf dynamics of a deciduous forest canopy: no response to elevated CO<Subscript>2</Subscript>

Leaf area index (LAI) and its seasonal dynamics are key determinants of terrestrial productivity and, therefore, of the response of ecosystems to a rising atmospheric CO₂ concentration. Despite the central importance of LAI, there is very little evidence from which to assess how forest LAI will respond to increasing [CO₂]. We assessed LAI and related leaf indices of a closed-canopy deciduous forest for 4 years in 25-m-diameter plots that were exposed to ambient or elevated CO₂ (542 ppm) in a free-air CO₂ enrichment (FACE) experiment. LAI of this Liquidambar styraciflua (sweetgum) stand was about 6 and was relatively constant year-to-year, including the 2 years prior to the onset of CO₂ treatment. LAI throughout the 1999–2002 growing seasons was assessed through a combination of data on photosynthetically active radiation (PAR) transmittance, mass of litter collected in traps, and leaf mass per unit area (LMA). There was no effect of [CO₂] on any expression of leaf area, including peak LAI, average LAI, or leaf area duration. Canopy mass and LMA, however, were significantly increased by CO₂ enrichment. The hypothesized connection between light compensation point (LCP) and LAI was rejected because LCP was reduced by [CO₂] enrichment only in leaves under full sun, but not in shaded leaves. Data on PAR interception also permitted calculation of absorbed PAR (APAR) and light use efficiency (LUE), which are key parameters connecting satellite assessments of terrestrial productivity with ecosystem models of future productivity. There was no effect of [CO₂] on APAR, and the observed increase in net primary productivity in elevated [CO₂] was ascribed to an increase in LUE, which ranged from 1.4 to 2.4 g MJ^–1. The current evidence seems convincing that LAI of non-expanding forest stands will not be different in a future CO₂-enriched atmosphere and that increases in LUE and productivity in elevated [CO₂] are driven primarily by functional responses rather than by structural changes. Ecosystem or regional models that incorporate feedbacks on resource use through LAI should not assume that LAI will increase with CO₂ enrichment of the atmosphere.     

Rising atmospheric carbon dioxide concentration and the future of C4 crops for food and fuel

Crops with the C₄ photosynthetic pathway are vital to global food supply, particularly in the tropical regions where human well-being and agricultural productivity are most closely linked. While rising atmospheric [CO₂] is the driving force behind the greater temperatures and water stress, which threaten to reduce future crop yields, it also has the potential to directly benefit crop physiology. The nature of C₄ plant responses to elevated [CO₂] has been controversial. Recent evidence from free-air CO₂ enrichment (FACE) experiments suggests that elevated [CO₂] does not directly stimulate C₄ photosynthesis. Nonetheless, drought stress can be ameliorated at elevated [CO₂] as a result of lower stomatal conductance and greater intercellular [CO₂]. Therefore, unlike C₃ crops for which there is a direct enhancement of photosynthesis by elevated [CO₂], C₄ crops will only benefit from elevated [CO₂] in times and places of drought stress. Current projections of future crop yields have assumed that rising [CO₂] will directly enhance photosynthesis in all situations and, therefore, are likely to be overly optimistic. Additional experiments are needed to evaluate the extent to which amelioration of drought stress by elevated [CO₂] will improve C₄ crop yields for food and fuel over the range of C₄ crop growing conditions and genotypes.     

Effects of [CO2] and nitrogen fertilization on soils planted with ponderosa pine

The effects of six years treatment with elevated [CO₂] (350, 525, and 700 μl l^-1) and nitrogen (N) (0, 10, and 20 g N m^-2 yr^-1) on soils, soil solution, and CO₂ efflux in an open-top chamber study with ponderosa pine (Pinus ponderosa Laws.) are described. The clearest [CO₂] effect was in year 6, when a pattern of lower soil N concentration and higher C/N ratio with elevated [CO₂] emerged. Statistically significant effects of elevated [CO₂] on soil total C, extractable P, exchangeable Mg²⁺, exchangeable Ca²⁺, base saturation, and soil solution HCO₃ ^- and NO₃ ^- were also found in various treatment combinations and at various times; however, these effects were inconsistent among treatments and years, and in many cases (P, Mg²⁺, Ca²⁺, base saturation) reflected pre-treatment differences. The use of homogenized buried soil bags did not improve the power to detect changes in soil C and N or help resolve the inconsistencies in soil C patterns. Nitrogen fertilization had the expected negative effects on exchangeable Ca²⁺, K⁺, and Mg²⁺ in year 6, presumably because of increased NO₃ ^- leaching, but had no consistent effect on soil C, N, or extractable P. This revised version was published online in June 2006 with corrections to the Cover Date.     

Impact of elevated CO2 concentration on ultrastructure of pericarp and composition of grain in three Triticum species of different ploidy levels

Triticum species of three ploidies were grown under ambient (375 μl/l) or elevated (FACE, 550 μl/l) CO₂ concentration [CO₂] to evaluate their response to CO₂ enrichment. The consistent effect of elevated CO₂ was an increase in concentration of starch and decrease in concentration of protein in the grain. Transmission electron micrographs revealed an increase in width and area of chloroplasts, and change in shape from elliptical in ambient to round in elevated [CO₂]. There was a corresponding increase in starch grain size and number in chloroplasts. The large starch grains distributed among thylakoids resulted in separation and distortion of internal membrane system in chloroplasts. The level of response was different in species of different ploidy levels. Maximum increase in starch concentration, and least decrease in protein concentration, was observed in Triticum dicoccoides, which also proved the most suitable species in terms of C:N ratio.     

Independent, Interactive, and Species-Specific Responses of Leaf Litter Decomposition to Elevated CO2 and O3 in a Northern Hardwood Forest

The future capacity of forest ecosystems to sequester atmospheric carbon is likely to be influenced by CO₂-mediated shifts in nutrient cycling through changes in litter chemistry, and by interactions with pollutants like O₃. We evaluated the independent and interactive effects of elevated CO₂ (560 μl l⁻¹) and O₃ (55 nl l l⁻¹) on leaf litter decomposition in trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) at the Aspen free air CO₂ enrichment (FACE) site (Wisconsin, USA). Fumigation treatments consisted of replicated ambient, +CO₂, +O₃, and +CO₂ + O₃ FACE rings. We followed mass loss and litter chemistry over 23 months, using reciprocally transplanted litterbags to separate substrate quality from environment effects. Aspen decayed more slowly than birch across all treatment conditions, and changes in decomposition dynamics of both species were driven by shifts in substrate quality rather than by fumigation environment. Aspen litter produced under elevated CO₂ decayed more slowly than litter produced under ambient CO₂, and this effect was exacerbated by elevated O₃. Similarly, birch litter produced under elevated CO₂ also decayed more slowly than litter produced under ambient CO₂. In contrast to results for aspen, however, elevated O₃ accelerated birch decay under ambient CO₂, but decelerated decay under enriched CO₂. Changes in decomposition rates (k-values) were due to CO₂- and O₃-mediated shifts in litter quality, particularly levels of carbohydrates, nitrogen, and tannins. These results suggest that in early-successional forests of the future, elevated concentrations of CO₂ will likely reduce leaf litter decomposition, although the magnitude of effect will vary among species and in response to interactions with tropospheric O₃.     

Contrasting Effects of Carbon Dioxide and Irradiance on the Acclimation of Photosynthesis in Developing Soybean Leaves

Leaves developed at high irradiance (I) often have higher photosynthetic capacity than those developed at low I, while leaves developed at elevated CO₂ concentration [CO₂] often have reduced photosynthetic capacity compared with leaves developed at lower [CO₂]. Because both high I and elevated [CO₂] stimulate photosynthesis of developing leaves, their contrasting effects on photosynthetic capacity at maturity suggest that the extra photosynthate may be utilized differently depending on whether I or [CO₂] stimulates photosynthesis. These experiments were designed to test whether relationships between photosynthetic income and the net accumulation of soluble protein in developing leaves, or relationships between soluble protein and photosynthetic capacity at full expansion differed depending on whether I or [CO₂] was varied during leaf development. Soybean plants were grown initially with a photosynthetic photon flux density (PPFD) of 950 µmol m^–2 s^–1 and 350 µmol [CO₂] mol^–1, then exposed to [CO₂] ranging from 135 to 1400 µmol mol^–1 for the last 3 d of expansion of third trifoliolate leaves. These results were compared with experiments in which I was varied at a constant [CO₂] of 350 µmol mol^–1 over the same developmental period. Increases in area and dry mass over the 3 d were determined along with daily photosynthesis and respiration. Photosynthetic CO₂ exchange characteristics and soluble protein content of leaves were determined at the end of the treatment periods. The increase in leaflet mass was about 28 % of the dry mass income from photosynthesis minus respiration, regardless of whether [CO₂] or I was varied, except that very low I or [CO₂] increased this percentage. Leaflet soluble protein per unit of area at full expansion had the same positive linear relationship to photosynthetic income whether [CO₂] or I was varied. For variation in I, photosynthetic capacity varied directly with soluble protein per unit area. This was not the case for variation in [CO₂]. Increasing [CO₂] reduced photosynthetic capacity per unit of soluble protein by up to a factor of 2.5, and photosynthetic capacity exhibited an optimum with respect to growth [CO₂]. Thus CO₂ did not alter the relationship between photosynthetic income and the utilization of photosynthate in the net accumulation of soluble protein, but did alter the relationship between soluble protein content and photosynthetic characteristics in this species.     

Atmospheric carbon dioxide and fertilizer nitrogen effects on radiation interception by rice 1

In order to predict the potential impacts of global change, it is important to understand the impact of increasing global atmospheric [CO₂] on the growth and yield of crop plants. The objectives of this study were to determine the interaction of N fertilization rates and atmospheric [CO₂] on radiation interception and radiation-use efficiency of rice (Oryza sativa L. cv. IR72) grown under tropical field conditions. Rice plants were grown inside open top chambers in a lowland rice field at the International Rice Research Institute in the Philippines at ambient (about 350 μmol mol^-1) or elevated (about 600 μmol mol^-1 during the 1993 wet season and 700 μmol mol^-1 during the 1994 dry season) in combination with three levels of applied N (0, 50 or 100 kg N ha^-1 in the wet season; 0, 90 or 200 kg N ha-1 in the dry season). Light interception was not directly affected by [CO₂], but elevated [CO₂] indirectly increased light interception through increasing total absorbed N. Plant N requirement for radiation interception was similar for rice grown under ambient [CO₂] or elevated [CO₂] treatments. The conversion efficiency of intercepted radiation to dry matter, radiation-use efficiency (RUE), was about 35% greater at elevated [CO₂] than at ambient [CO₂]. The relationship between leaf N and RUE was curvilinear. At ambient [CO₂], RUE was fairly stable across levels of leaf N, but leaf N less than about 2.5% resulted in lower RUE for plants grown with elevated [CO₂] than for plant grown at ambient [CO₂]. Decreased leaf N with increased [CO₂], therefore decreased RUE of rice plants grown at elevated [CO₂]. When predicting responses of rice to elevated [CO₂], RUE should be adjusted with a decrease in leaf N. This revised version was published online in June 2006 with corrections to the Cover Date.     

Wood properties of Populus and Betula in long‐term exposure to elevated CO2 and O3

We studied the interactive effects of elevated concentrations of CO₂ and O₃ on radial growth and wood properties of four trembling aspen (Populus tremuloides Michx.) clones and paper birch (Betula papyrifera Marsh.) saplings. The material for the study was collected from the Aspen FACE (free‐air CO₂ enrichment) experiment in Rhinelander (WI, USA). Trees had been exposed to four treatments [control, elevated CO₂ (560 ppm), elevated O₃ (1.5 times ambient) and combined CO₂ + O₃] during growing seasons 1998–2008. Most treatment responses were observed in the early phase of experiment. Our results show that the CO₂‐ and O₃‐exposed aspen trees displayed a differential balance between efficiency and safety of water transport. Under elevated CO₂, radial growth was enhanced and the trees had fewer but hydraulically more efficient larger diameter vessels. In contrast, elevated O₃ decreased radial growth and the diameters of vessels and fibres. Clone‐specific decrease in wood density and cell wall thickness was observed under elevated CO₂. In birch, the treatments had no major impacts on wood anatomy or wood density. Our study indicates that short‐term impact studies conducted with young seedlings may not give a realistic view of long‐term ecosystem responses.