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.     

Free Air CO2 Enrichment of potato (Solanum tuberosum L.): development, growth and yield

A FACE (Free Air CO₂ Enrichment) experiment was carried out on Potato (Solanum tuberosum L., cv. Primura) in 1995 in Italy. Three FACE rings were used to fumigate circular field plots of 8 m diameter while two rings were used as controls at ambient CO₂ concentrations. Four CO₂ exposure levels were used in the rings (ambient, 460, 560 and 660 μmol mol^–1). Phenology and crop development, canopy surface temperature, above- and below-ground biomass were monitored during the growing season. Crop phenology was affected by elevated CO₂, as the date of flowering was progressively anticipated in the 660, 560, 460 μmol mol^–1 treatments. Crop development was not affected significantly as plant height, leaf area and the number of leaves per plant were the same in the four treatments. Elevated atmospheric CO₂ levels had, instead, a significant effect on the accumulation of total nonstructural carbohydrates (TNC = soluble sugars + starch) in the leaves during a sunny day. Specific leaf area was decreased under elevated CO₂ with a response that paralleled that of TNC concentrations. This reflected the occurrence of a progressive increase of photosynthetic rates and carbon assimilation in plants exposed to increasingly higher levels of atmospheric CO₂. Tuber growth and final tuber yield were also stimulated by rising CO₂ levels. When calculated by regression of tuber yield vs. the imposed levels of CO₂concentration, yield stimulation was as large as 10% every 100 μmol mol^–1 increase, which translated into over 40% enhancement in yield under 660 μmol mol^–1. This was related to a higher number of tubers rather than greater mean tuber mass or size. Leaf senescence was accelerated under elevated CO₂ and a linear relationship was found between atmospheric CO₂ levels and leaf reflectance measured at 0.55 μm wavelength. We conclude that significant CO₂ stimulation of yield has to be expected for potato under future climate scenarios, and that crop phenology will be affected as well.     

The turnover of carbon pools contributing to soil CO2 and soil respiration in a temperate forest exposed to elevated CO2 concentration

Soil carbon is returned to the atmosphere through the process of soil respiration, which represents one of the largest fluxes in the terrestrial C cycle. The effects of climate change on the components of soil respiration can affect the sink or source capacity of ecosystems for atmospheric carbon, but no current techniques can unambiguously separate soil respiration into its components. Long‐term free air CO₂ enrichment (FACE) experiments provide a unique opportunity to study soil C dynamics because the CO₂ used for fumigation has a distinct isotopic signature and serves as a continuous label at the ecosystem level. We used the ¹³C tracer at the Duke Forest FACE site to follow the disappearance of C fixed before fumigation began in 1996 (pretreatment C) from soil CO₂ and soil‐respired CO₂, as an index of belowground C dynamics during the first 8 years of the experiment. The decay of pretreatment C as detected in the isotopic composition of soil‐respired CO₂ and soil CO₂ at 15, 30, 70, and 200 cm soil depth was best described by a model having one to three exponential pools within the soil system. The majority of soil‐respired CO₂ (71%) originated in soil C pools with a turnover time of about 35 days. About 55%, 50%, and 68% of soil CO₂ at 15, 30, and 70 cm, respectively, originated in soil pools with turnover times of less than 1 year. The rest of soil CO₂ and soil‐respired CO₂ originated in soil pools that turn over at decadal time scales. Our results suggest that a large fraction of the C returned to the atmosphere through soil respiration results from dynamic soil C pools that cannot be easily detected in traditionally defined soil organic matter standing stocks. Fast oxidation of labile C substrates may prevent increases in soil C accumulation in forests exposed to elevated [CO₂] and may consequently result in shorter ecosystem C residence times.     

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.     

Productivity and water use of wheat under free-air CO2 enrichment

A free-air CO₂ enrichment (FACE) experiment was conducted at Maricopa, Arizona, on wheat from December 1992 through May 1993. The FACE apparatus maintained the CO₂ concentration, [CO₂], at 550 μmol mol^?1 across four replicate 25-m-diameter circular plots under natural conditions in an open field. Four matching Control plots at ambient [CO₂] (about 370 μmol mol^?1) were also installed in the field. In addition to the two levels of [CO₂], there were ample (Wet) and limiting (Dry) levels of water supplied through a subsurface drip irrigation system in a strip, split-plot design. Measurements were made of net radiation, R_n; soil heat flux, G_o; soil temperature; foliage or surface temperature; air dry and wet bulb temperatures; and wind speed. Sensible heat flux, H, was calculated from the wind and temperature measurements. Latent heat flux, λET, and evapotranspiration, ET, were determined as the residual in the energy balance. The FACE treatment reduced daily total R_n by an average 4%. Daily FACE sensible heat flux, H, was higher in the FACE plots. Daily latent heat flux, λET, and evapotranspiration, ET, were consistently lower in the FACE plots than in the Control plots for most of the growing season, about 8% on the average. Net canopy photosynthesis was stimulated by an average 19 and 44% in the Wet and Dry plots, respectively, by elevated [CO₂] for most of the growing season. No significant acclimation or down regulation was observed. There was little above-ground growth response to elevated [CO₂] early in the season when temperatures were cool. Then, as temperatures warmed into spring, the FACE plants grew about 20% more than the Control plants at ambient [CO₂], as shown by above-ground biomass accumulation. Root biomass accumulation was also stimulated about 20%. In May the FACE plants matured and senesced about a week earlier than the Controls in the Wet plots. The FACE plants averaged 0.6 °C warmer than the Controls from February through April in the well-watered plots, and we speculate that this temperature rise contributed to the earlier maturity. Because of the acceleration of senescence, there was a shortening of the duration of grain filling, and consequently, there was a narrowing of the final biomass and yield differences. The 20% mid-season growth advantage of FACE shrunk to about an 8% yield advantage in the Wet plots, while the yield differences between FACE and Control remained at about 20% in the Dry plots.     

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.     

Effects of chronic ozone and elevated atmospheric CO2 concentrations on ribulose-1,5-bisphosphate in soybean (Glycine max)

Ribulose-1,5-bisphosphate (RuBP) pool size was determined at regular intervals during the growing season to understand the effects of tropospheric ozone concentrations, elevated atmospheric carbon dioxide concentrations and their interactions on the photosynthetic limitation by RuBP regeneration. Soybean (Glycine max [L.] Merr. cv. Essex) was grown from seed to maturity in open-top field chambers in charcoal-filtered air (CF) either without (22 nmol O₃ mol^?1) or with added O₃ (83 nmol mol^?1) at ambient (AA, 369 μmol CO₂ mol^?1) or elevated CO₂ (710 μmol mol^?1). The RuBP pool size generally declined with plant age in all treatments when expressed on a unit leaf area and in all treatments but CF-AA when expressed per unit ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco; EC 4.1.1.39) binding site. Although O₃ in ambient CO₂ generally reduced the RuBP pool per unit leaf area, it did not change the RuBP pool per unit Rubisco binding site. Elevated CO₂, in CF or O₃-fumigated air, generally had no significant effect on RuBP pool size, thus mitigating the negative O₃ effect. The RuBP pools were below 2 mol mol^?1 binding site in all treatments for most of the season, indicating limiting RuBP regeneration capacity. These low RuBP pools resulted in increased RuBP regeneration via faster RuBP turnover, but only in CF air and during vegetative and flowering stages at elevated CO₂. Also, the low RuBP pool sizes did not always reflect RuBP consumption rates or the RuBP regeneration limitation relative to potential carboxylation (%RuBP). Rather, %RuBP increased linearly with decrease in the RuBP pool turnover time. These data suggest that amelioration of damage from O₃ by elevated atmospheric CO₂ to the RuBP regeneration may be in response to changes in the Rubisco carboxylation.     

Direct and indirect effects of elevated CO2 on leaf respiration in a forest ecosystem

We measured the short‐term direct and long‐term indirect effects of elevated CO₂ on leaf dark respiration of loblolly pine (Pinus taeda) and sweetgum (Liquidambar styraciflua) in an intact forest ecosystem. Trees were exposed to ambient or ambient + 200 µmol mol^?1 atmospheric CO₂ using free‐air carbon dioxide enrichment (FACE) technology. After correcting for measurement artefacts, a short‐term 200 µmol mol^?1 increase in CO₂ reduced leaf respiration by 7–14% for sweetgum and had essentially no effect on loblolly pine. This direct suppression of respiration was independent of the CO₂ concentration under which the trees were grown. Growth under elevated CO₂ did not appear to have any long‐term indirect effects on leaf maintenance respiration rates or the response of respiration to changes in temperature (Q₁₀, R₀). Also, we found no relationship between mass‐based respiration rates and leaf total nitrogen concentrations. Leaf construction costs were unaffected by growth CO₂ concentration, although leaf construction respiration decreased at elevated CO₂ in both species for leaves at the top of the canopy. We conclude that elevated CO₂ has little effect on leaf tissue respiration, and that the influence of elevated CO₂ on plant respiratory carbon flux is primarily through increased biomass.     

Is photosynthetic acclimation to free-air CO<Subscript>2</Subscript> enrichment (FACE) related to a strong competition for the assimilatory power between carbon assimilation and nitrogen assimilation in rice leaf?

Net photosynthetic rate (P _N) of leaves grown under free-air CO₂ enriched condition (FACE, about 200 μmol mol⁻¹ above ambient air) was significantly lower than P _N of leaves grown at ambient CO₂ concentration (AC) when measured at CO₂ concentration of 580 μmol mol⁻¹. This difference was found in rice plants grown at normal nitrogen supply (25 g m⁻²; NN-plants) but not in plants grown at low nitrogen supply (15 g m⁻²; LN-plants). Namely, photosynthetic acclimation to FACE was observed in NN-plants but not in LN-plants. Different from the above results measured in a period of continuous sunny days, such photosynthetic acclimation occurred in NN-plants, however, it was also observed in LN-plants when P _N was measured before noon of the first sunny day after rain. Hence strong competition for the assimilatory power between nitrogen (N) and carbon (C) assimilations induced by an excessive N supply may lead to the photosynthetic acclimation to FACE in NN-plants. The hypothesis is supported by the following facts: FACE induced significant decrease in both apparent photosynthetic quantum yield (Φ_c) and ribulose-1,5-bisphosphate (RuBP) content in NN-plants but not in LN-plants.     

A free-air enrichment system for exposing tall forest vegetation to elevated atmospheric CO2

A free-air CO₂ enrichment (FACE) system was designed to permit the experimental exposure of tall vegetation such as stands of forest trees to elevated atmospheric CO₂ concentrations ([CO₂]_a) without enclosures that alter tree microenvironment. We describe a prototype FACE system currently in operation in forest plots in a maturing loblolly pine (Pinus taeda L.) stand in North Carolina, USA. The system uses feedback control technology to control [CO₂] in a 26 m diameter forest plot that is over 10 m tall, while monitoring the 3D plot volume to characterize the whole-stand CO₂ regime achieved during enrichment. In the second summer season of operation of the FACE system, atmospheric CO₂ enrichment was conducted in the forest during all daylight hours for 96.7% of the scheduled running time from 23 May to 14 October with a preset target [CO₂] of 550 μmol mol^–1, ≈ 200 μmol mol^–1 above ambient [CO₂]. The system provided spatial and temporal control of [CO₂] similar to that reported for open-top chambers over trees, but without enclosing the vegetation. The daily average daytime [CO₂] within the upper forest canopy at the centre of the FACE plot was 552 ± 9 μmol mol^–1 (mean ± SD). The FACE system maintained 1-minute average [CO₂] to within ± 110 μmol mol^–1 of the target [CO₂] for 92% of the operating time. Deviations of [CO₂] outside of this range were short-lived (most lasting < 60 s) and rare, with fewer than 4 excursion events of a minute or longer per day. Acceptable spatial control of [CO₂] by the system was achieved, with over 90% of the entire canopy volume within ± 10% of the target [CO₂] over the exposure season. CO₂ consumption by the FACE system was much higher than for open-top chambers on an absolute basis, but similar to that of open-top chambers and branch bag chambers on a per unit volume basis. CO₂ consumption by the FACE system was strongly related to windspeed, averaging 50 g CO₂ m^–3 h^–1 for the stand for an average windspeed of 1.5 m s^–1 during summer. The [CO₂] control results show that the free-air approach is a tractable way to study long-term and short-term alterations in trace gases, even within entire tall forest ecosystems. The FACE approach permits the study of a wide range of forest stand and ecosystem processes under manipulated [CO₂]_a that were previously impossible or intractable to study in true forest ecosystems.     

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.     

Acclimation response of spring wheat in a free-air CO2 enrichment (FACE) atmosphere with variable soil nitrogen regimes. 2. Net assimilation and stomatal conductance of leaves

Atmospheric CO₂ concentration continues to rise. It is important, therefore, to determine what acclimatory changes will occur within the photosynthetic apparatus of wheat (Triticum aestivum L. cv. Yecora Rojo) grown in a future high-CO₂ world at ample and limited soil N contents. Wheat was grown in an open field exposed to the CO₂ concentration of ambient air [370 μmol (CO₂) mol⁻¹; Control] and air enriched to ∼200 μmol (CO₂) mol⁻¹ above ambient using a Free-Air CO₂ Enrichment (FACE) apparatus (main plot). A High (35 g m⁻²) or Low (7 and 1.5 g m⁻² for 1996 and 1997, respectfully) level of N was applied to each half of the main CO₂ treatment plots (split-plot). Under High-N, FACE reduced stomatal conductance (g _s) by 30% at mid-morning (2 h prior to solar noon), 36% at midday (solar noon) and 27% at mid-afternoon (2.5 h after solar noon), whereas under Low-N, g _s was reduced by as much as 31% at mid-morning, 44% at midday and 28% at mid-afternoon compared with Control. But, no significant CO₂ × N interaction effects occurred. Across seasons and growth stages, daily accumulation of carbon (A′) was 27% greater in FACE than Control. High-N increased A′ by 18% compared with Low-N. In contrast to results for g _s, however, significant CO₂ × N interaction effects occurred because FACE increased A′ by 30% at High-N, but by only 23% at Low-N. FACE enhanced the seasonal accumulation of carbon (A′′) by 29% during 1996 (moderate N-stress), but by only 21% during 1997 (severe N-stress). These results support the premise that in a future high-CO₂ world an acclimatory (down-regulation) response in the photosynthetic apparatus of field-grown wheat is anticipated. They also demonstrate, however, that the stimulatory effect of a rise in atmospheric CO₂ on carbon gain in wheat can be maintained if nutrients such as nitrogen are in ample supply. This revised version was published online in June 2006 with corrections to the Cover Date.     

Arbuscular mycorrhizal fungi benefit from 7 years of free air CO2 enrichment in well-fertilized grass and legume monocultures

Rising atmospheric carbon dioxide partial pressure (pCO₂) and nitrogen (N) deposition are important components of global environmental change. In the Swiss free air carbon dioxide enrichment (FACE) experiment, the effect of altered atmospheric pCO₂ (35 vs. 60 Pa) and the influence of two different N‐fertilization regimes (14 vs. 56 g N m^?2 a^?1) on root colonization by arbuscular mycorrhizal fungi (AMF) and other fungi (non‐AMF) of Lolium perenne and Trifolium repens were studied. Plants were grown in permanent monoculture plots, and fumigated during the growth period for 7 years. At elevated pCO₂ AMF and non‐AMF root colonization was generally increased in both plant species, with significant effects on colonization intensity and on hyphal and non‐AMF colonization. The CO₂ effect on arbuscules was marginally significant (P=0.076). Moreover, the number of small AMF spores (≤100 μm) in the soils of monocultures (at low‐N fertilization) of both plant species was significantly increased, whereas that of large spores (>100 μm) was increased only in L. perenne plots. N fertilization resulted in a significant decrease of root colonization in L. perenne, including the AMF parameters, hyphae, arbuscules, vesicles and intensity, but not in T. repens. This phenomenon was probably caused by different C‐sink limitations of grass and legume. Lacking effects of CO₂ fumigation on intraradical AMF structures (under high‐N fertilization) and no response to N fertilization of arbuscules, vesicles and colonization intensity suggest that the function of AMF in T. repens was non‐nutritional. In L. perenne, however, AM symbiosis may have amended N nutrition, because all root colonization parameters were significantly increased under low‐N fertilization, whereas under high‐N fertilization only vesicle colonization was increased. Commonly observed P‐nutritional benefits from AMF appeared to be absent under the phosphorus‐rich soil conditions of our field experiment. We hypothesize that in well‐fertilized agricultural ecosystems, grasses benefit from improved N nutrition and legumes benefit from increased protection against pathogens and/or herbivores. This is different from what is expected in nutritionally limited plant communities.     

Combined effects of elevated CO2 and air temperature on carbon assimilation of Pinus taeda trees

Branches of 22-year-old loblolly pine (Pinus taeda, L.) trees growing in a plantation were exposed to ambient CO₂, ambient + 165 μmol mol^?1 CO₂ or ambient + 330 μmol mol^?1 CO₂ concentrations in combination with ambient or ambient + 2°C air temperatures for 3 years. Field measurements in the third year indicated that net carbon assimilation was enhanced in the elevated CO₂ treatments in all seasons. On the basis of A/C_i, curves, there was no indication of photosynthetic down-regulation. Branch growth and leaf area also increased significantly in the elevated CO₂ treatments. The imposed 2°C increase in air temperature only had slight effects on net assimilation and growth. Compared with the ambient CO₂ treatment, rates of net assimilation were ～1·6 times greater in the ambient + 165 μmol mol^?1 CO₂ treatment and 2·2 times greater in the ambient + 330 μmol mol^?1 CO₂ treatment. These ratios did not change appreciably in measurements made in all four seasons even though mean ambient air temperatures during the measurement periods ranged from 12·6 to 28·2°C. This indicated that the effect of elevated CO₂ concentrations on net assimilation under field conditions was primarily additive. The results also indicated that the effect of elevated CO₂ (+ 165 or + 330 μmol mol^?1) was much greater than the effect of a 2°C increase in air temperature on net assimilation and growth in this species.     

Elevated atmospheric [CO2] stimulates sugar accumulation and cellulose degradation rates of rice straw

Rice straw can serve as potential material for bioenergy production. However, the quantitative effects of increasing atmospheric carbon dioxide concentration [CO₂] on rice straw quality and the resulting consequences for bioenergy utilization are largely unknown. In this study, two rice varieties, WYJ and LY, that have been shown previously to have a weak and strong stimulatory response to rising [CO₂], respectively, were grown with and without additional CO₂ at China free‐air carbon dioxide enrichment (FACE) platform. Qualitative and quantitative measurements in response to [CO₂] included straw biomass (including leaf, sheath, and stem), the concentration of nonstructural and structural carbohydrates, the syringyl‐to‐guaiacyl (S/G) ratio of lignin, glucose and xylose release from structural carbohydrate, total sugar release by enzymatic saccharification, and sugar yield and the ratio of cellulose and hemicellulose degradation. Elevated [CO₂] significantly increased straw biomass and nonstructural carbohydrate contents while enhancing the degraded ratio of structural carbohydrates as indicated by the decreased lignin content and increased S/G ratio. Overall, total sugar yield (g m^?2) in rice straw significantly increased by 27.1 and 57% for WYJ and LY at elevated [CO₂], respectively. These findings, while preliminary, suggest that rice straw quality and potential biofuel utilization may improve as a function of rising [CO₂].     

The effects of free-air CO2 enrichment and soil water availability on spatial and seasonal patterns of wheat root growth

Spring wheat [ Triticum aestivum (L). cv. Yecora Rojo] was grown from December 1992 to May 1993 under two atmospheric CO₂ concentrations, 550 μmol mol^–1 for high-CO₂ plots, and 370 μmol mol^–1 for control plots, using a Free-Air CO₂ Enrichment (FACE) apparatus. In addition to the two levels of atmospheric CO₂, there were ample and limiting levels of water supply through a subsurface trip irrigation system in a strip, split-plot design. In order to examine the temporal and spatial root distribution, root cores were extracted at six growth stages during the season at in-row and inter-row positions using a soil core device (86 mm ID, 1.0 m length). Such information would help determine whether and to what extent root morphology is changed by alteration of two important factors, atmospheric CO₂ and soil water, in this agricultural ecosystem. Wheat root growth increased under elevated CO₂ conditions during all observed developmental stages. A maximum of 37% increase in total root dry mass in the FACE vs. Control plots was observed during the period of stem elongation. Greater root growth rates were calculated due to CO₂ enhancement until anthesis. During the early vegetative growth, root dry mass of the inter-row space was significantly higher for FACE than for Control treatments suggesting that elevated CO₂ promoted the production of first-order lateral roots per main axis. Then, during the reproductive period of growth, more branching of lateral roots in the FACE treatment occurred due to water stress. Significant higher root dry mass was measured in the inter-row space of the FACE plots where soil water supply was limiting. These sequential responses in root growth and morphology to elevated CO₂ and reduced soil water supports the hypothesis that plants grown in a high-CO₂ environment may better compensate soil-water-stress conditions.     

Above- and below-ground responses of C3–C4 species mixtures to elevated CO2 and soil water availability

We evaluated the influences of CO₂[Control, ～ 370 µ mol mol ^{? 1}; 200 µ mol mol ^{? 1} above ambient applied by free‐air CO₂ enrichment (FACE)] and soil water (Wet, Dry) on above‐ and below‐ground responses of C₃ (cotton, Gossypium hirsutum) and C₄ (sorghum, Sorghum bicolor) plants in monocultures and two density mixtures. In monocultures, CO₂ enrichment increased height, leaf area, above‐ground biomass and reproductive output of cotton, but not sorghum, and was independent of soil water treatment. In mixtures, cotton, but not sorghum, above‐ground biomass and height were generally reduced compared to monocultures, across both CO₂ and soil water treatments. Density did not affect individual plant responses of either cotton or sorghum across the other treatments. Total (cotton + sorghum) leaf area and above‐ground biomass in low‐density mixtures were similar between CO₂ treatments, but increased by 17–21% with FACE in high‐density mixtures, due to a 121% enhancement of cotton leaf area and a 276% increase in biomass under the FACE treatment. Total root biomass in the upper 1.2 m of the soil was not influenced by CO₂ or by soil water in monoculture or mixtures; however, under dry conditions we observed significantly more roots at lower soil depths ( > 45 cm). Sorghum roots comprised 81–85% of the total roots in the low‐density mixture and 58–73% in the high‐density mixture. CO₂‐enrichment partly offset negative effects of interspecific competition on cotton in both low‐ and high‐density mixtures by increasing above‐ground biomass, with a greater relative increase in the high‐density mixture. As a consequence, CO₂‐enrichment increased total above‐ground yield of the mixture at high density. Individual plant responses to CO₂ enrichment in global change models that evaluate mixed plant communities should be adjusted to incorporate feedbacks for interspecific competition. Future field studies in natural ecosystems should address the role that a CO₂‐mediated increase in C₃ growth may have on subsequent vegetation change.     

Carbon-based secondary and structural compounds in Mediterranean shrubs growing near a natural CO2 spring

We studied carbon‐based secondary and structural compounds (CBSSCs) in Myrtus communis, Erica arborea, and Juniperus communis co‐occurring in a natural CO₂ spring site and in a nearby control site in a Mediterranean environment. Leaf concentrations of phenolics and CBSSCs, such as lignin, cellulose, and hemicellulose, total nonstructural carbohydrates (TNCs), and lipids were measured monthly (phenolics) and every two months (the other compounds) throughout a year. There was a slight seasonal trend towards maximum concentrations of most of these CBSSCs during autumn–winter and minimum values during the spring season, particularly in Myrtus communis. For most of the CBSSCs and species, there were no consistent or significant patterns in response to the elevated [CO₂] (c. 700 μmol mol^?1) of the spring site. These results were not due to a dilution effect by increased structural or nonstructural carbon. Therefore, in contrast to many experimental studies of CO₂ enrichment, mainly conducted for short periods, there were no greater concentrations of phenolics, and, as in many of these studies, there were neither greater concentrations of the other CBSSCs. These results do not agree with the predictions of the carbon source‐sink hypotheses. Possible causes of this disagreement are discussed. These causes include the complex heterogeneous environmental conditions and the variability of resource availabilities in the field, photosynthetic down‐regulation, and/or the homeostatic and evolutionary nature of organisms. These results suggest evolutionary adaptive responses to changes in CO₂. They also suggest caution in attributing increased CBSSC concentrations to elevated [CO₂] at long‐term scale in natural conditions, and therefore in their implications for plant–herbivore interactions and for decomposition.     

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.     

Net soil carbon input under ambient and elevated CO2 concentrations: isotopic evidence after 4 years

Elevation of atmospheric CO₂ concentration is predicted to increase net primary production, which could lead to additional C sequestration in terrestrial ecosystems. Soil C input was determined under ambient and Free Atmospheric Carbon dioxide Enrichment (FACE) conditions for Lolium perenne L. and Trifolium repens L. grown for four years in a sandy‐loam soil. The ¹³C content of the soil organic matter C had been increased by 5‰ compared to the native soil by prior cropping to corn (Zea mays) for > 20 years. Both species received low or high amounts of N fertilizer in separate plots. The total accumulated above‐ground biomass produced by L. perenne during the 4‐year period was strongly dependent on the amount of N fertilizer applied but did not respond to increased CO₂. In contrast, the total accumulated above‐ground biomass of T. repens doubled under elevated CO₂ but remained independent of N fertilizer rate. The C:N ratio of above‐ground biomass for both species increased under elevated CO₂ whereas only the C:N ratio of L. perenne roots increased under elevated CO₂. Root biomass of L. perenne doubled under elevated CO₂ and again under high N fertilization. Total soil C was unaffected by CO₂ treatment but dependent on species. After 4 years and for both crops, the fraction of new C (F‐value) under ambient conditions was higher (P= 0.076) than under FACE conditions: 0.43 vs. 0.38. Soil under L. perenne showed an increase in total soil organic matter whereas N fertilization or elevated CO₂ had no effect on total soil organic matter content for both systems. The net amount of C sequestered in 4 years was unaffected by the CO₂ concentration (overall average of 8.5 g C kg^?1 soil). There was a significant species effect and more new C was sequestered under highly fertilized L. perenne. The amount of new C sequestered in the soil was primarily dependent on plant species and the response of root biomass to CO₂ and N fertilization. Therefore, in this FACE study net soil C sequestration was largely depended on how the species responded to N rather than to elevated CO₂.