Decades-long effects of high CO<Subscript>2</Subscript> concentration on soil nitrogen dynamics at a natural CO<Subscript>2</Subscript> spring

The effects of high atmospheric CO₂ concentration ([CO₂]) on ecosystem processes have been explored using temporal facilities such as open-top-chambers and free-air CO₂ enrichment. However, the effects of high [CO₂] on soil properties takes decades and may not be captured by short-term experiments. Natural CO₂ springs provide a unique opportunity to study the long-term effects of high [CO₂]. In this study, we investigated soil properties at a natural CO₂ spring. We found that the amounts of total carbon (C) and nitrogen (N) stored in the soil at the high [CO₂] site exceeded those in the reference site by 60 and 30%, respectively. The effects of high [CO₂] were large in the upper slope position where the canopy openness was high and plants grew faster, but no effects were detected in the lowest position where the canopy openness was lower (half of that at the upper slope position). In contrast, effects of high [CO₂] on soil N dynamics, such as N mineralization and nitrification rates, did not exhibit a slope gradient. This suggests that effects of high [CO₂] differed among soil stoichiometric characteristics and N dynamics. These complicated effects of high [CO₂] imply that the future effects of high [CO₂] on ecosystems could vary widely in conjunction with environmental conditions such as light availability and/or topographic conditions.     

Autumnal leaf abscission of sugar maple is not delayed by atmospheric CO<Subscript>2</Subscript> enrichment

To investigate the effects of atmospheric CO₂ enrichment on physiology and autumnal leaf phenology, we exposed 3-year-old sugar maple (Acer saccharum Marsh.) seedlings to 800 (A8), 600 (A6), and 400 μL(CO₂) L^–1 (AA) in nine continuous stirred tank reactor (CSTR) chambers during the growing season of 2014. Leaf abscission timing, abscised leaf area percentages, leaf number, light-saturated net photosynthetic rate (P_Nmax), leaf area, accumulative growth rates, and biomass were determined and assessed. The results suggested the following: (1) no significant differences were found in the timing of leaf abscission in the three CO₂-concentration treatments; (2) P_Nmax was continuously stimulated to the greatest extent in A8 at 319% and 160% in A6 until the end of the growing season, respectively; and (3) leaf number, leaf area, and accumulative height growth all significantly increased by elevated CO₂, which led to a 323% increase in A8 biomass and 235% in A6 biomass after 156-d fumigation. In summary, the results suggest, the timing of leaf abscission of sugar maple in fall was not modified by CO₂ enrichment, the increased carbon gain by elevated CO₂ was mainly due to increased leaf area, more leaves, and the continuously enhanced high photosynthesis throughout the growing season instead of the leaf life span.     

Photosynthetic CO<Subscript>2</Subscript> uptake in seedlings of two tropical tree species exposed to oscillating elevated concentrations of CO<Subscript>2</Subscript>

Do short-term fluctuations in CO₂ concentrations at elevated CO₂ levels affect net CO₂ uptake rates of plants? When exposed to 600 μl CO₂ l^?1, net CO₂ uptake rates in shoots or leaves of seedlings of two tropical C₃ tree species, teak (Tectona grandis L. f.) and barrigon [Pseudobombax septenatum (Jacq.) Dug.], increased by 28 and 52% respectively. In the presence of oscillations with half-cycles of 20 s, amplitude of ca. 170 μl CO₂ l^?1 and mean of 600 μl CO₂ l^?1, the stimulation in net CO₂ uptake by the two species was reduced to 19 and 36%, respectively, i.e. the CO₂ stimulation in photosynthesis associated with a change in exposure from 370 to 600 μl CO₂ l^?1 was reduced by a third in both species. Similar reductions in CO₂-stimulated net CO₂ uptake were observed in T. grandis exposed to 40-s oscillations. Rates of CO₂ efflux in the dark by whole shoots of T. grandis decreased by 4.8% upon exposure of plants grown at 370 μl CO₂ l^?1 to 600 μl CO₂ l^?1. The potential implications of the observations on CO₂ oscillations and dark respiration are discussed in the context of free-air CO₂ enrichment (FACE) systems in which short-term fluctuations of CO₂ concentration are a common feature.     

The effects of elevated atmospheric CO<Subscript>2</Subscript> and nitrogen amendments on subsurface CO<Subscript>2</Subscript> production and concentration dynamics in a maturing pine forest

Profiles of subsurface soil CO₂ concentration, soil temperature, and soil moisture, and throughfall were measured continuously during the years 2005 and 2006 in 16 locations at the free air CO₂ enrichment facility situated within a temperate loblolly pine (Pinus taeda L.) stand. Sampling at these locations followed a 4 by 4 replicated experimental design comprised of two atmospheric CO₂ concentration levels (ambient [CO₂]^a, ambient + 200 ppmv, [CO₂]^e) and two soil nitrogen (N) deposition levels (ambient, ambient + fertilization at 11.2 g_N m⁻² year⁻¹). The combination of these measurements permitted indirect estimation of belowground CO₂ production and flux profiles in the mineral soil. Adjacent to the soil CO₂ profiles, direct (chamber-based) measurements of CO₂ fluxes from the soil–litter complex were simultaneously conducted using the automated carbon efflux system. Based on the measured soil CO₂ profiles, neither [CO₂]^e nor N fertilization had a statistically significant effect on seasonal soil CO₂, CO₂ production, and effluxes from the mineral soil over the study period. Soil moisture and temperature had different effects on CO₂ concentration depending on the depth. Variations in CO₂ were mostly explained by soil temperature at deeper soil layers, while water content was an important driver at the surface (within the first 10 cm), where CO₂ pulses were induced by rainfall events. The soil effluxes were equal to the CO₂ production for most of the time, suggesting that the site reached near steady-state conditions. The fluxes estimated from the CO₂ profiles were highly correlated to the direct measurements when the soil was neither very dry nor very wet. This suggests that a better parameterization of the soil CO₂ diffusivity is required for these soil moisture extremes.     

Modeling soil CO<Subscript>2</Subscript> emissions from ecosystems

We present a new soil respiration model, describe a formal model testing procedure, and compare our model with five alternative models using an extensive data set of observed soil respiration. Gas flux data from rangeland soils that included a large number of measurements at low temperatures were used to model soil CO₂ emissions as a function of soil temperature and water content. Our arctangent temperature function predicts that Q₁₀ values vary inversely with temperature and that CO₂ fluxes are significant below 0 °C. Independent data representing a broad range of ecosystems and temperature values were used for model testing. The effects of plant phenology, differences in substrate availability among sites, and water limitation were accounted for so that the temperature equations could be fairly evaluated. Four of the six tested models did equally well at simulating the observed soil CO₂ respiration rates. However, the arctangent variable Q₁₀ model agreed closely with observed Q₁₀ values over a wide range of temperatures (r² = 0.94) and was superior to published variable Q₁₀ equations using the Akaike information criterion (AIC). The arctangent temperature equation explained 16–85% of the observed intra-site variability in CO₂ flux rates. Including a water stress factor yielded a stronger correlation than temperature alone only in the dryland soils. The observed change in Q₁₀ with increasing temperature was the same for data sets that included only heterotrophic respiration and data sets that included both heterotrophic and autotrophic respiration.     

Carbon allocation in shoots of alpine treeline conifers in a CO<Subscript>2</Subscript> enriched environment

With a new approach we assessed the relative contribution of stored and current carbon compounds to new shoot growth in alpine treeline conifers. Within a free air CO₂ enrichment experiment at the alpine treeline in Switzerland, ¹³C-depleted fossil CO₂ was used to trace new carbon in the two tree species Larix decidua L. and Pinus uncinata Ramond over two subsequent years. The deciduous L. decidua was found to supply new shoot growth (structural woody part) by 46% from storage. Surprisingly, the evergreen P. uncinata, assumed to use current-year photosynthates, also utilized a considerable fraction of storage (42%) for new wood growth. In contrast, the needles of P. uncinata were built up almost completely from current-year photosynthates. The isotopic composition of different wood carbon fractions revealed a similar relative allocation of current and stored assimilates to various carbon fractions. Elevated CO₂ influenced the composition of woody tissue in a species-specific way, e.g. the water soluble fraction decreased in pine in 2001 but increased in larch in 2002 compared to ambient CO₂. Heavy defoliation applied as an additional treatment factor in the second year of the experiment decreased the lipophilic fraction in current-year wood in both species compared to undefoliated trees. We conclude that storage may play an important role for new shoot growth in these treeline conifers and that altered carbon availability (elevated CO₂, defoliation) results in significant changes in the relative amount of mobile carbon fractions in woody tissue. In particular, stored carbon seems to be of greater importance in the evergreen P. uncinata than has been previously thought.     

Microgravity does not alter plant stand gas exchange of wheat at moderate light levels and saturating CO<Subscript>2</Subscript> concentration

Plant stand gas exchange was measured nondestructively in microgravity during the Photosynthesis Experiment Subsystem Testing and Operations experiment conducted onboard the International Space Station. Rates of evapotranspiration and photosynthesis measured in space were compared with ground controls to determine if microgravity directly affects whole-stand gas exchange of Triticum aestivum. During six 21-day experiment cycles, evapotranspiration was determined continuously from water addition rates to the nutrient delivery system, and photosynthesis was determined from the amount of CO₂ added to maintain the chamber CO₂ concentration setpoint. Plant stand evapotranspiration, net photosynthesis, and water use efficiency were not altered by microgravity. Although leaf area was significantly reduced in microgravity-grown plants compared to ground control plants, leaf area distribution was not affected enough to cause significant differences in the amounts of light absorbed by the flight and ground control plant stands. Microgravity also did not affect the response of evapotranspiration to changes in chamber vapor pressure difference of 12-day-old wheat plant stands. These results suggest that gravity naïve plants grown at moderate light levels (300 mol m^–2 s^–1) behave the same as ground control plants. This implies that future plant-based regenerative life support systems can be sized using 1 g data because water purification and food production rates operate at nearly the same rates as in 1 g at moderate light levels. However, it remains to be verified whether the present results are reproducible in plants grown under stronger light levels.     

Long-term effects of elevated atmospheric CO<Subscript>2</Subscript> on species composition and productivity of a southern African C<Subscript>4</Subscript> dominated grassland in the vicinity of a CO<Subscript>2</Subscript> exhalation

We describe the long-term effects of a CO₂ exhalation, created more than 70 years ago, on a natural C₄ dominated sub-tropical grassland in terms of ecosystem structure and functioning. We tested whether long-term CO₂ enrichment changes the competitive balance between plants with C₃ and C₄ photosynthetic pathways and how CO₂ enrichment has affected species composition, plant growth responses, leaf properties and soil nutrient, carbon and water dynamics. Long-term effects of elevated CO₂ on plant community composition and system processes in this sub-tropical grassland indicate very subtle changes in ecosystem functioning and no changes in species composition and dominance which could be ascribed to elevated CO₂ alone. Species compositional data and soil δ¹³C isotopic evidence suggest no detectable effect of CO₂ enrichment on C₃:C₄ plant mixtures and individual species dominance. Contrary to many general predictions C₃ grasses did not become more abundant and C₃ shrubs and trees did not invade the site. No season length stimulation of plant growth was found even after 5 years of exposure to CO₂ concentrations averaging 610 μmol mol⁻¹. Leaf properties such as total N decreased in the C₃ but not C₄ grass under elevated CO₂ while total non-structural carbohydrate accumulation was not affected. Elevated CO₂ possibly lead to increased end-of-season soil water contents and this result agrees with earlier studies despite the topographic water gradient being a confounding problem at our research site. Long-term CO₂ enrichment also had little effect on soil carbon storage with no detectable changes in soil organic matter found. There were indications that potential soil respiration and N mineralization rates could be higher in soils close to the CO₂ source. The conservative response of this grassland suggests that many of the reported effects of elevated CO₂ on similar ecosystems could be short duration experimental artefacts that disappear under long-term elevated CO₂ conditions.     

Experimental evidence that terrestrial carbon subsidies increase CO<Subscript>2</Subscript> flux from lake ecosystems

Subsidies are donor-controlled inputs of nutrients and energy that can affect ecosystem-level processes in a recipient environment. Lake ecosystems receive large inputs of terrestrial carbon (C) in the form of dissolved organic matter (DOM). DOM inputs may energetically subsidize heterotrophic bacteria and determine whether lakes function as sources or sinks of atmospheric CO₂. I experimentally tested this hypothesis using a series of mesocosm experiments in New England lakes. In the first experiment, I observed that CO₂ flux increased by 160% 4 days following a 1,000 m C addition in the form of DOM. However, this response was relatively short lived, as there was no effect of DOM enrichment on CO₂ flux beyond 8 days. In a second experiment, I demonstrated that peak CO₂ flux from mesocosms in two lakes increased linearly over a broad DOM gradient (slope for both lakes=0.02±0.001 mm CO₂·m^–2 day^–1 per m DOC, mean±SE). Concomitant changes in bacterial productivity and dissolved oxygen strengthen the inference that increasing CO₂ flux resulted from the metabolism of DOM. I conducted two additional studies to test whether DOM-correlated attributes were responsible for the observed change in plankton metabolism along the subsidy gradient. First, terrestrial DOM reduced light transmittance, but experimental shading revealed that this was not responsible for the observed patterns of CO₂ flux. Second, organically bound nitrogen (N) and phosphorus (P) accompanied DOM inputs, but experimental nutrient additions (without organic C) caused mesocosms to be satuated with CO₂. Together, these results suggest that C content of terrestrial DOM may be an important subsidy for freshwater bacteria that can influence whether recipient aquatic ecosystems are sources or sinks of atmospheric CO₂.     

The effects of elevated atmospheric [CO<Subscript>2</Subscript>] on Norway spruce needle parameters

Studies of selected morphological needle parameters were carried out on young (17–19 year old) Norway spruce trees cultivated inside glass domes at ambient (A, 370 μmol (CO₂) mol⁻¹) and elevated (E, 700 μmol (CO₂) mol⁻¹) atmospheric CO₂ concentrations [CO₂] beginning in 1997. Annual analyses performed from 2002 to 2004 revealed higher values for needle length (especially for current needles, up to 18%) and projected needle area (up to 13%) accompanied by lower values for specific needle area (up to 15% lower, as quantified by needle mass to projected area ratio) in the E treatment compared to the A treatment. Statistically significant differences for most of the investigated morphological parameters were found in young needles in the well irradiated sun-adapted crown parts, particularly under water-limiting soil conditions in 2003. This was likely a result of different water relations in E compared to A trees as investigated under temperate water stress (Kuper et al. in Biol Plantarum 50:603–609, 2006). Furthermore, E trees had much higher absorbing root area, which modified and enhanced root:shoot as well as root:conductive stem area proportions. These hydraulic properties and early seasonal stimulation of photosynthesis forced advanced needle development in E trees, particularly under limited soil water conditions. The number of needles per unit shoot length was found to be unaffected by elevated [CO₂].     

Root-derived CO<Subscript>2</Subscript> flux from a tropical peatland

Tropical peatlands release significant quantities of greenhouse gases to the atmosphere, yet the relative contributions of heterotrophic and autotrophic respiration to net CO₂ fluxes remains sparsely quantified. We used a combination of in situ trenching and vegetation removal in ex situ pots to quantify root-derived CO₂ under two plant functional types within a mixed species forest. Trenching significantly reduced surface CO₂ flux, indicating that approximately two-thirds of the released CO₂ was derived from roots. In contrast, ex situ vegetation removal in pots indicated that root-derived CO₂ accounted for 27% of the total CO₂ flux for Campnosperma panamensis, a broadleaved evergreen tree, and 49% for Raphia taedigera, a canopy palm. The results show that root-derived CO₂ is a major contribution to net CO₂ emissions in tropical peatlands, and that the magnitude of the emissions is affected by plant species composition. This is important in the context of land use change driving alterations in vegetation cover.     

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.     

Elevated CO<Subscript>2</Subscript> increased photosynthesis and yield without decreasing stomatal conductance in broomcorn millet

Broomcorn millet (Panicum miliaceum L.) is one of the important C₄ crops in the semiarid regions of northern China. It is a close relative of biofuel crop switchgrass. Yet, there is no information on how these crops might respond to a climate change in China. In order to gain insight into such a response, we studied the effect of elevated CO₂ concentration (EC) on broomcorn millet. The changes in leaf photosynthesis, chlorophyll fluorescence, morphological parameters, biomass and yield in response to EC [i.e., + 200 µmol(CO₂) mol^?1] over two years were determined at the open-top chamber (OTC) experimental facility in north China. EC increased net photosynthetic rate, stomatal conductance, intercellular CO₂ concentration, transpiration rate, instantaneous transpiration efficiency, effective quantum yield of PSII photochemistry, and photochemical quenching coefficient of fully expanded flag leaves. Maximal quantum yield of PSII photochemistry declined under EC in 2013, but was not affected in 2014. EC significantly decreased intrinsic efficiency of PSII in 2013, but increased in 2014. Leaf nonphotochemical quenching decreased under EC both in 2013 and 2014. EC significantly enhanced the aboveground biomass and yield by average of 31.4 and 25.5% in both years, respectively. The increased yield of broomcorn millet under EC occurred due to the enhanced number of grains per plant. We concluded that photosynthesis of broomcorn millets was improved through increased stomatal conductance in leaves under EC, which led to an increase in height, stem diameter, aboveground biomass, and yield. This study extends our understanding of the response of this ancient C₄ crop to elevated CO₂ concentration.     

CO<Subscript>2</Subscript> sequestration in plants: lesson from divergent strategies

Most organisms inhabiting earth feed directly or indirectly on the products synthesized by the reaction of photosynthesis, which at the current atmospheric CO₂ levels operates only at two thirds of its peak efficiency. Restricting the photorespiratory loss of carbon and thereby improving the efficiency of photosynthesis is seen by many as a good option to enhance productivity of food crops. Research during last half a century has shown that several plant species developed CO₂-concentrating mechanism (CCM) to restrict photorespiration under lower concentration of available CO₂. CCMs are now known to be operative in several terrestrial and aquatic plants, ranging from most advanced higher plants to algae, cyanobacteria and diatoms. Plants with C₄ pathway of photosynthesis (where four-carbon compound is the first product of photosynthesis) or crassulacean acid metabolism (CAM) may consistently operate CCM. Some plants however can undergo a shift in photosynthetic metabolism only with change in environmental variables. More recently, a shift in plant photosynthetic metabolism is reported at high altitude where improved efficiency of CO₂ uptake is related to the recapture of photorespiratory loss of carbon. Of the divergent CO₂ assimilation strategies operative in different oraganisms, the capacity to recapture photorespiratory CO₂ could be an important approach to develop plants with efficient photosynthetic capacity.     

Effects of elevated atmospheric CO<Subscript>2</Subscript> on the nutritional ecology of C<Subscript>3</Subscript> and C<Subscript>4</Subscript> grass-feeding caterpillars

It is plausible that the nutritional quality of C₃ plants will decline more under elevated atmospheric CO₂ than will the nutritional quality of C₄ plants, causing herbivorous insects to increase their feeding on C₃ plants relative to C₄ plants. We tested this hypothesis with a C₃ and C₄ grass and two caterpillar species with different diet breadths. Lolium multiflorum (C₃) and Bouteloua curtipendula (C₄) were grown in outdoor open top chambers at ambient (370 ppm) or elevated (740 ppm) CO₂. Bioassays compared the performance and digestive efficiencies of Pseudaletia unipuncta (a grass-specialist noctuid) and Spodoptera frugiperda (a generalist noctuid). As expected, the nutritional quality of L. multiflorum changed to a greater extent than did that of B. curtipendula when grown in elevated CO₂; levels of protein (considered growth limiting) declined in the C₃ grass, while levels of carbohydrates (sugar, starch and fructan) increased. However, neither insect species increased its feeding rate on the C₃ grass to compensate for its lower nutritional quality when grown in an elevated CO₂ atmosphere. Consumption rates of P. unipuncta and S. frugiperda were higher on the C₃ grass than the C₄ grass, the opposite of the result expected for a compensatory response to the lower nutritional quality of the C₄ grass. Although our results do not support the hypothesis that grass-specialist insects compensate for lower nutritional quality by increasing their consumption rates more than do generalist insects, the performance of the specialist was greater than that of the generalist on each grass species and at both CO₂ levels. Mechanisms other than compensatory feeding, such as increased nutrient assimilation efficiency, appear to determine the relative performance of these herbivores. Our results also provide further evidence against the hypothesis that C₄ grasses would be avoided by insect herbivores because a large fraction of their nutrients is unavailable to herbivores. Instead, our results are consistent with the hypothesis that C₄ grasses are poorer host plants primarily because of their lower nutrient levels, higher fiber levels, and greater toughness.     

Hydrothermal Stability of Adenine Under Controlled Fugacities of N<Subscript>2</Subscript>, CO<Subscript>2</Subscript> and H<Subscript>2</Subscript>

An experimental study has been carried out on the stability of adenine (one of the five nucleic acid bases) under hydrothermal conditions. The experiments were performed in sealed autoclaves at 300 degrees C under fugacities of CO(2), N(2) and H(2) supposedly representative of those in marine hydrothermal systems on the early Earth. The composition of the gas phase was obtained from the degradation of oxalic acid, sodium nitrite and ammonium chloride, and the oxidation of metallic iron. The results of the experiments indicate that after 200 h, adenine is still present in detectable concentration in the aqueous phase. In fact, the concentration of adenine does not seem to be decreasing after approximately 24 h, which suggests that an equilibrium state may have been established with the inorganic constituents of the hydrothermal fluid. Such a conclusion is corroborated by independent thermodynamic calculations.     

Effect of high CO<Subscript>2</Subscript> concentrations on the growth and macromolecular composition of a heat- and high-light-tolerant microalga

A green microalga, Acutodesmus sp., a close relative of Acutodesmus deserticola, was isolated from the wastewater discharges of an oil refinery in India. This study examined the effects of light intensity, temperature, pH, and high-CO₂ treatments (up to 20 %) on the growth of the alga and investigated the effects of different CO₂ treatments on its macromolecular composition (protein, carbohydrate, and lipids). Under controlled laboratory conditions, the alga showed high growth rates over a wide range of light (up to 700 μmol photons m^?2 s^?1), temperature (up to 40 °C), and pH (5–10) conditions. In the stationary phase, the highest protein and carbohydrate content was found to be 71.52 and 40.72 % of dry weight at 5 and 15 % CO₂, respectively. After 5 days of cultivation, the maximum dry weight biomass attained in these cultures was 1.149, 1.99, 1.75, and 1.65 g L^?1 at 5, 10, 15, and 20 % CO₂, respectively, indicating that this strain has significant tolerance to CO₂. These results indicate that this strain is a promising candidate for use in biofixation of CO₂ from the flue gases emitted by industries, and it also has a strong potential as a feedstock for value-added substances.     

CO<Subscript>2</Subscript> enrichment increases element concentrations in grass mixtures by changing species abundances

Atmospheric carbon dioxide (CO₂) enrichment may increase plant growth more than the uptake of chemical elements from soil. Increased CO₂ also may alter element levels in biomass from multi-species vegetation by changing plant species abundances. We measured concentrations of ten elements in aboveground tissues of three C₄ grasses that had been exposed for 2–3 growing seasons to a continuous gradient in CO₂ from 250 to 500 μmol mol⁻¹. The grasses, Bouteloua curtipendula, Schizachyrium scoparium, and Sorghastrum nutans, are competitive dominants in assemblages of tallgrass prairie vegetation growing on each of three soil types along a field CO₂ gradient in central Texas, USA. Our objective was to determine whether CO₂ influences element concentrations in grass mixtures by changing concentrations in individual species or shifting species abundances. Increased CO₂ had little effect on element concentrations in grasses compared to differences observed among grass species and soils. Increasing CO₂ from the pre-Industrial to elevated levels reduced the phosphorus concentration in grasses grown on a clay and sandy loam soil. Concentrations of most other elements did not respond to CO₂ treatment. Cover of the mid-grass Bouteloua declined at higher CO₂ levels as cover of the taller grass Sorghastrum increased. Concentrations of several elements were lower in Bouteloua than Sorghastrum; hence, this exchange of species at higher CO₂ increased element concentrations in grass assemblages. Potential consequences include an improvement in the nutritional quality of plants for herbivores. Results highlight the underappreciated impact that CO₂ enrichment may have on ecosystem functioning by changing plant composition.     

Effect of elevated CO<Subscript>2</Subscript> levels on leaf starch,nitrogen and photosynthesis of plants growing at three natural CO<Subscript>2</Subscript> springs in Japan

Plant communities around natural CO₂ springs have been exposed to elevated CO₂ levels over many generations and give us a unique opportunity to investigate the effects of long-term elevated CO₂ levels on wild plants. We searched for natural CO₂ springs in cool temperate climate regions in Japan and found three springs that were suitable for studying long-term responses of plants to elevated levels of CO₂: Ryuzin-numa, Yuno-kawa and Nyuu. At these CO₂ springs, the surrounding air was at high CO₂ concentration with no toxic gas emissions throughout the growth season, and there was natural vegetation around the springs. At each site, high-CO₂ (HC) and low-CO₂ (LC) plots were established, and three dominant species at the shrub layers were used for physiological analyses. Although the microenvironments were different among the springs, dicotyledonous species growing at the HC plots tended to have more starch and less nitrogen per unit dry mass in the leaves than those growing at the LC plots. In contrast, monocotyledonous species growing in the HC and LC plots had similar starch and nitrogen concentrations. Photosynthetic rates at the mean growth CO₂ concentration were higher in HC plants than LC plants, but photosynthetic rates at a common CO₂ concentration were lower in HC plants. Efficiency of water and nitrogen use of leaves at growth CO₂ concentration was greatly increased in HC plants. These results suggest that natural plants growing in elevated CO₂ levels under cool temperate climate conditions have down-regulated their photosynthetic capacity but that they increased photosynthetic rates and resource use efficiencies due to the direct effect of elevated CO₂ concentration.     

Leaf senescence in response to elevated atmospheric CO<Subscript>2</Subscript> concentration and low nitrogen supply

This review reports the physiological and metabolic changes in plants during development under elevated atmospheric carbon dioxide concentration and/or limited-nitrogen supply in order to establish their effects on leaf senescence induction. Elevated CO₂ concentration and nitrogen supply modify gene expression, protein content and composition, various aspects of photosynthesis, sugar metabolism, nitrogen metabolism, and redox state in plants. Elevated CO₂ usually causes sugar accumulation and decreased nitrogen content in plant leaves, leading to imbalanced C/N ratio in mature leaves, which is one of the main factors behind premature senescence in leaves. Elevated CO₂ and low nitrogen decrease activities of some antioxidant enzymes and thus increase H₂O₂ production. These changes lead to oxidative stress that results in the degradation of photosynthetic pigments and eventually induce senescence. However, this accelerated leaf senescence under conditions of elevated CO₂ and limited nitrogen can mobilize nutrients to growing organs and thus ensure their functionality.