Assessment of grain quality in terms of functional group response to elevated [CO2], water,and nitrogen using a meta‐analysis: Grain protein,zinc, and iron under future climate

The increasing [CO₂] in the atmosphere increases crop productivity. However, grain quality of cereals and pulses are substantially decreased and consequently compromise human health. Meta‐analysis techniques were employed to investigate the effect of elevated [CO₂] (e[CO₂]) on protein, zinc (Zn), and iron (Fe) concentrations of major food crops (542 experimental observations from 135 studies) including wheat, rice, soybean, field peas, and corn considering different levels of water and nitrogen (N). Each crop, except soybean, had decreased protein, Zn, and Fe concentrations when grown at e[CO₂] concentration (≥550 μmol/mol) compared to ambient [CO₂] (a[CO₂]) concentration (≤380 μmol/mol). Grain protein, Zn, and Fe concentrations were reduced under e[CO₂]; however, the responses of protein, Zn, and Fe concentrations to e[CO₂] were modified by water stress and N. There was an increase in Fe concentration in soybean under medium N and wet conditions but nonsignificant. The reductions in protein concentrations for wheat and rice were ~5%–10%, and the reductions in Zn and Fe concentrations were ~3%–12%. For soybean, there was a small and nonsignificant increase of 0.37% in its protein concentration under medium N and dry water, while Zn and Fe concentrations were reduced by ~2%–5%. The protein concentration of field peas decreased by 1.7%, and the reductions in Zn and Fe concentrations were ~4%–10%. The reductions in protein, Zn, and Fe concentrations of corn were ~5%–10%. Bias in the dataset was assessed using a regression test and rank correlation. The analysis indicated that there are medium levels of bias within published meta‐analysis studies of crops responses to free‐air [CO₂] enrichment (FACE). However, the integration of the influence of reporting bias did not affect the significance or the direction of the [CO₂] effects.     

Effect of elevated atmospheric CO2 and vegetation type on microbiota associated with decomposing straw

Straw from wheat plants grown at ambient and elevated atmospheric CO₂ concentrations was placed in litterbags in a grass fallow field and a wheat field. The CO₂ treatment induced an increase in straw concentration of ash‐free dry mass from 84% to 93% and a decrease in nitrogen concentration from 0.43% to 0.34%. After five months of decomposition, less than 50% of the straw was decomposed. The content of ash‐free dry mass remaining in straw from plants grown at elevated CO₂ was significantly higher than that from plants grown at ambient CO₂ (4.02 vs. 3.69 g AFDM per litterbag in the fallow field and 3.40 vs. 2.67 g AFDM per litterbag when buried in the wheat field). The immobilization of nitrogen during decomposition was significantly higher in the ambient straw, and there was a significant negative correlation between the content of organic matter remaining per litterbag and the nitrogen concentration in the recovered straw samples. After five months of decomposition, hyphal biomass was significantly lower in straw from plants grown at elevated CO₂ (? 30% and ?13% in the fallow and wheat field, respectively). Bacterial biomass was not significantly affected by the CO₂ induced changes in the litter quality, but the lower decomposition rate and fewer bacterial grazers in the straw from plants grown at elevated CO₂ together indicate reduced microbial activity and turnover. Notwithstanding this, these data show that growth at elevated atmospheric CO₂ concentration results in slower decomposition of wheat straw, but the effect is probably of minor importance compared to the effect of varying crops, agricultural practise or changing land use.     

The effects of enhanced ozone and enhanced carbon dioxide concentrations on biomass, pigments and antioxidative enzymes in spruce needles (Picea abies L.)

During one growing period, 5-year-old spruce trees (Picea abies L., Karst.) were exposed in environmental chambers to elevated concentrations of carbon dioxide (750 cm³ m^?3) and ozone (008 cm³ m^?3) as single variables or in combination. Control concentrations of the gases were 350cm³ m^?3CO₂ and 0.02 cm³ m ^?3 ozone. To investigate whether an elevated CO₂ concentration can prevent adverse ozone effects by reducing oxidative stress, the activities of the protective enzymes superoxide dismutase, catalase and peroxidase were determined. Furthermore, shoot biomass, pigment and protein contents of two needle age classes were investigated. Ozone caused pigment reduction and visible injury in the previous year's needles and growth reduction in the current year's shoots. In the presence of elevated concentrations of ozone and CO₂, growth reduction in the current year's shoots was prevented, but emergence of visible damage in the previous year's needles was only delayed and pigment reduction was still found. Elevated concentrations of ozone or CO₂ as single variables caused a significant reduction in the activities of superoxide dismutase and catalase in the current year's needles. Minimum activities of superoxide dismutase and catalase and decreased peroxidase activities were found in both needle age classes from spruce trees grown at enhanced concentrations of both CO₂ and ozone. These results suggest a reduced tolerance to oxidative stress in spruce trees under conditions of elevated concentrations of both CO₂ and ozone.     

Direct and acclimatory responses of stomatal conductance to elevated carbon dioxide in four herbaceous crop species in the field

In order to separate the net effect of growth at elevated [CO₂] on stomatal conductance (g_s) into direct and acclimatory responses, mid‐day values of g_s were measured for plants grown in field plots in open‐topped chambers at the current ambient [CO₂], which averaged 350 μmol mol^?1 in the daytime, and at ambient + 350 μmol mol^?1[CO₂] for winter wheat, winter barley, potato and sorghum. The acclimatory response was determined by comparing g_s measured at 700 μmol mol^?1[CO₂] for plants grown at the two [CO₂]. The direct effect of increasing [CO₂] from 350 to 700 μmol mol^?1 was determined for plants grown at the lower concentration. Photosynthetic rates were measured concurrently with g_s. For all species, growth at the higher [CO₂] significantly reduced g_s measured at 700 μmol mol^?1[CO₂]. The reduction in g_s caused by growth at the higher [CO₂] was larger for all species on days with low leaf to air water vapour pressure difference for a given temperature, which coincided with highest conductances and also the smallest direct effects of increased [CO₂] on conductance. For barley, there was no other evidence for stomatal acclimation, despite consistent down‐regulation of photosynthetic rate in plants grown at the higher [CO₂]. In wheat and potato, in addition to the vapour pressure difference interaction, the magnitude of stomatal acclimation varied directly in proportion to the magnitude of down‐regulation of photosynthetic rate through the season. In sorghum, g_s consistently exhibited acclimation, but there was no down‐regulation of photosynthetic rate. In none of the species except barley was the direct effect the larger component of the net reduction in g_s when averaged over measurement dates. The net effect of growth at elevated [CO₂] on mid‐day g_s resulted from unique combinations of direct and acclimatory responses in the various species.     

Interaction of Elevated Ultraviolet-B Radiation and CO(2) on Productivity and Photosynthetic Characteristics in Wheat, Rice, and Soybean

Wheat (Triticum aestivum L. cv Bannock), rice (Oryza sativa L. cv IR-36), and soybean (Glycine max [L.] Merr cv Essex) were grown in a factorial greenhouse experiment to determine if CO₂-induced increases in photosynthesis, biomass, and yield are modified by increases in ultraviolet (UV)-B radiation corresponding to stratospheric ozone depletion. The experimental conditions simulated were: (a) an increase in CO₂ concentration from 350 to 650 microliters per liter; (b) an increase in UV-B radiation corresponding to a 10% ozone depletion at the equator; and (c) a and b in combination. Seed yield and total biomass increased significantly with elevated CO₂ in all three species when compared to the control. However, with concurrent increases in UV-B and CO₂, no increase in either seed yield (wheat and rice) or total biomass (rice) was observed with respect to the control. In contrast, CO₂-induced increases in seed yield and total plant biomass were maintained or increased in soybean within the elevated CO₂, UV-B environment. Whole leaf gas exchange indicated a significant increase in photosynthesis, apparent quantum efficiency (AQE) and water-use-efficiency (WUE) with elevated CO₂ in all 3 species. Including elevated UV-B radiation with high CO₂ eliminated the effect of high CO₂ on photosynthesis and WUE in rice and the increase in AQE associated with high CO₂ in all species. Elevated CO₂ did not change the apparent carboxylation efficiency (ACE) in the three species although the combination of elevated CO₂ and UV-B reduced ACE in wheat and rice. The results of this experiment illustrate that increased UV-B radiation may modify CO₂-induced increases in biomass, seed yield and photosynthetic parameters and suggest that available data may not adequately characterize the potential effect of future, simultaneous changes in CO₂ concentration and UV-B radiation.     

Interacting effects of CO2 concentration, temperature and nitrogen supply on the photosynthesis and composition of winter wheat leaves

Winter wheat (Triticum aestivum L., cv. Mercia) was grown at two different atmospheric CO₂ concentrations (350 and 700 μmol mol⁻¹), two temperatures [ambient temperature (i.e. tracking the open air) and ambient +4°C] and two rates of nitrogen supply (equivalent to 489 kg ha⁻¹ and 87 kg ha⁻¹). Leaves grown at 700 μmol mol⁻¹ CO₂ had slightly greater photosynthetic capacity (10% mean increase over the experiment) than those grown at ambient CO₂ concentration, but there were no differences in carboxylation efficiency or apparent quantum yield. The amounts of chlorophyll, soluble protein and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) per unit leaf area did not change with long-term exposure to elevated CO₂ concentration. Thus winter wheat, grown under simulated field conditions, for which total biomass was large compared to normal field production, did not experience loss of components of the photosynthetic system or loss of photosynthetic competence with elevated CO₂ concentration. However, nitrogen supply and temperature had large effects on photosynthetic characteristics but did not interact with elevated CO₂ concentration. Nitrogen deficiency resulted in decreases in the contents of protein, including Rubisco, and chlorophyll, and decreased photosynthetic capacity and carboxylation efficiency. An increase in temperature also reduced these components and shortened the effective life of the leaves, reducing the duration of high photosynthetic capacity.     

Effect of CO2 enrichment on non-structural carbohydrates in leaves, stems and ears of spring wheat

Field-grown spring wheat (Triticum aestivum L. cv. Dragon) was exposed to ambient and elevated CO₂ concentrations (1.5 and 2 times ambient) in open-top chambers. Contents of non-structural carbohydrates were analysed enzymatically in leaves, stems and ears six times during the growing season. The impact of elevated CO₂ on wheat carbohydrates was non-significant in most harvests. However, differences in the carbohydrate contents due to elevated CO₂ were found in all plant compartments. Before anthesis, at growth stage (GS) 30 (the stem is 1 cm to the shoot apex), the plants grown in elevated CO₂ contained significantly more water soluble carbohydrates (WSC), fructans, starch and total non-structural carbohydrates (TNC) in the leaves in comparison with the plants grown in ambient CO₂. It is hypothesised that the plants from the treatments with elevated CO₂ were sink-limited at GS30. After anthesis, the leaf WSC and TNC contents of the plants from elevated CO₂ started to decline earlier than those of the plants from ambient CO₂. This may indicate that the leaves of plants grown in the chambers with elevated CO₂ senesced earlier. Elevated CO₂ accelerated grain development: 2 weeks after anthesis, the plants grown in elevated CO₂ contained significantly more starch and significantly less fructans in the ears compared to the plants grown in ambient CO₂. Elevated CO₂ had no effect on ear starch and TNC contents at the final harvest. Increasing the CO₂ concentration from 360 to 520 μmol mol^?1 had a larger effect on wheat non-structural carbohydrates than the further increase from 520 to 680 μmol mol^?1. The results are discussed in relation to the effects of elevated CO₂ on yield and yield components.     

Effects of CO(2) Concentration on Rubisco Activity, Amount, and Photosynthesis in Soybean Leaves

Growth at an elevated CO₂ concentration resulted in an enhanced capacity for soybean (Glycine max L. Merr. cv Bragg) leaflet photosynthesis. Plants were grown from seed in outdoor controlled-environment chambers under natural solar irradiance. Photosynthetic rates, measured during the seed filling stage, were up to 150% greater with leaflets grown at 660 compared to 330 microliters of CO₂ per liter when measured across a range of intercellular CO₂ concentrations and irradiance. Soybean plants grown at elevated CO₂ concentrations had heavier pod weights per plant, 44% heavier with 660 compared to 330 microliters of CO₂ per liter grown plants, and also greater specific leaf weights. Ribulose 1,5-bisphosphate carboxylase/oxygenase (rubisco) activity showed no response (mean activity of 96 micromoles of CO₂ per square meter per second expressed on a leaflet area basis) to short-term (~1 hour) exposures to a range of CO₂ concentrations (110-880 microliters per liter), nor was a response of activity (mean activity of 1.01 micromoles of CO₂ per minute per milligram of protein) to growth CO₂ concentration (160-990 microliters per liter) observed. The amount of rubisco protein was constant, as growth CO₂ concentration was varied, and averaged 55% of the total leaflet soluble protein. Although CO₂ is required for activation of rubisco, results indicated that within the range of CO₂ concentrations used (110-990 microliters per liter), rubisco activity in soybean leaflets, in the light, was not regulated by CO₂.     

Effects of elevated CO2 on the foraging behavior of cotton bollworm, Helicoverpa armigera

Effects of elevated CO2 on the foraging behavior of cotton bollworm Helicoverpa arrnigera Hübner reared on milky grains of spring wheat grown in ambient, 550μL/L and 750μL/L CO2 concentration atmospheres in open-top chambers （OTC） were studied. The results indicated that： （i） elevated CO2 significantly affected both the type and amount of food eaten by H.arrnigera reared on milky grains of ambient CO2-grown wheat were significant higher than those for bollworm larvae reared on wheat grains grown in 550 and 750μL/L CO2 atmospheres; （ii） when bollworm larvae were reared on mixed milky grains from different CO2-grown wheat （food-choice condition）, larval duration increased significantly-pupal weight, adult longevity, and fecundity decreased significantly, comparing with those reared on milky grains of ambient CO2-grown wheat, 550μL/L CO2-grown wheat and 750μL/L CO2-grown wheat respectively; （iii） significant decreases in the contents of fructose and gross protein （GP） and significant increases in the contents of glucose, amylose, total saccharides （TSC）, TSC： GP ratio, free amino acids and soluble protein in the wheat grains with CO2 rising; （iv） and selected-foraging amount/food-choice index of cotton bollworm H.armigera were significantly positive correlated with the contents of fructose and GP of wheat grains, but they had significantly negative relationships with the contents of glucose, amylose, TSC and TSC： GP ratio of wheat grains.     

Effects of elevated CO2 concentration, supplemental irrigation and nitrogenous fertilizer application on rain-fed spring wheat yield

It is expected that the CO₂ concentration of the Earth’s atmosphere will reach 600–1000 ppm by the end of the 21st century. Therefore, in this study, we evaluated the effects of elevated CO₂ concentrations on the development of rain-fed spring wheat in an attempt to identify a practical pathway to increase crop production. To accomplish this, a field experiment was conducted at Guyuan Experimental Station in a semiarid region of China during 2005–2007. During this experiment, the CO₂ concentration was increased to 40.0 ppm and supplemental irrigation and nitrogenous fertilizer (N fertilizer) were applied. The experimental results showed that the elevated CO₂ concentration significantly improved the thousand-grain weight and the grain number per spike. Furthermore, supplemental irrigation and N fertilizer application during the elongation and booting stage of rain-fed spring wheat in conjunction with an elevated CO₂ concentration improved the water use efficiency (WUE), nitrogen use efficiency (NUE), thousand-grain weight, and the yield by 14.6%, 39.6%, 9.3%, and 14.7%, respectively, when compared to groups subjected to the same treatment but not grown under elevated CO₂ concentrations. Furthermore, the spring wheat yield was improved by 81.8% in response to an elevated CO₂ concentration, 60 mm of supplemental irrigation and applied N fertilizer (37.5 g m^?2 NH₄NO₃). However, the presence of an elevated CO₂ concentration without supplemental irrigation and N fertilizer only resulted in an increase in the wheat yield of 7.8%. Consequently, the combination of elevated CO₂ concentration, supplemental irrigation and N fertilizer application played an important role in the improvement of WUE, NUE, thousand-grain weight, and grain yield of rain-fed spring wheat in this region.     

Effects of free-air CO2 enrichment on the development of the photosynthetic apparatus in wheat, as indicated by changes in leaf proteins

A spring wheat crop was grown at ambient and elevated (550 μmol mol^?1) CO₂ concentrations under free-air CO₂ enrichment (FACE) in the field. Four experimental blocks, each comprising 21-m-diameter FACE and control experimental areas, were used. CO₂ elevation was maintained day and night from crop emergence to final grain harvest. This experiment provided a unique opportunity to examine the hypothesis that CO₂ elevation in the field would lead to acclimatory changes within the photosynthetic apparatus under open field conditions and lo assess whether acclimation was affected by crop developmental stage, leaf ontogeny and leaf age. Change in the photosynthetic apparatus was assessed by measuring changes in the composition of total leaf and thylakoid polypeptides separated by SDS-PAGE. For leaves at completion of emergence of the blade, growth at the elevated CO₂ concentration had no apparent effect on the amount of any of the major proteins of the photosynthetic apparatus regardless of the leaf examined. Leaf 5 on the main stem was in full sunlight at emergence, but then became shaded progressively as 3–4 further leaves formed above with continued development of the crop. By 35 d following completion of blade emergence, leaf 5 was in shade. At this point, the chlorophyll alb ratio had declined by 26% both in plants grown at the control CO₂ concentration and in those grown at the elevated CO₂ concentration, which is indicative of shade acclimation. The ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) content declined by 45% in the control leaves, but by 60% in the leaves grown at the elevated CO₂ concentration. The light- harvesting complex of photosystcm II (LHCII) and the chlorophyll content showed no decrease and no difference between treatments, indicating that the decrease in Rubisco was not an effect of earlier senescence in the leaves at the elevated CO₂ concentration. Following completion of the emergence of the flag-leaf blade, the elevated-CO₂ treatment inhibited the further accumulation of Rubisco which was apparent in control leaves over the subsequent 14 d. From this point onwards, the flag leaves from both treatments showed a loss of Rubisco, which was far more pronounced in the elevated-CO₂ treatment, so that by 36 d the Rubisco content of these leaves was just 70% of that of the controls and by 52 d it was only 20%. At 36 d, there was no decline in chlorophyll, LHCII or the chloroplast ATPase coupling factor (CFI) in the elevated CO₂ concentration treatment relative to the control. By 52 d, all of these proteins showed a significant decline relative to the control. This indicates that the decreased concentration of Rubisco at this final stage probably reflected earlier senescence in the elevated-CO₂ treatment, but that this was preceded by a CO₂-concentration-dependent decline in Rubisco.     

Effects of elevated CO2 and temperature-grown red and sugar maple on gypsy moth performance

Few studies have investigated how tree species grown under elevated CO₂ and elevated temperature alter the performance of leaf‐feeding insects. The indirect effects of an elevated CO₂ concentration and temperature on leaf phytochemistry, along with potential direct effects on insect growth and consumption, may independently or interactively affect insects. To investigate this, we bagged larvae of the gypsy moth on leaves of red and sugar maple growing in open‐top chambers in four CO₂/temperature treatment combinations: (i) ambient temperature, ambient CO₂; (ii) ambient temperature, elevated CO₂ (+ 300 μL L^?1 CO₂); (iii) elevated temperature (+ 3.5°C), ambient CO₂; and (iv) elevated temperature, elevated CO₂. For both tree species, leaves grown at elevated CO₂ concentration were significantly reduced in leaf nitrogen concentration and increased in C: N ratio, while neither temperature nor its interaction with CO₂ concentration had any effect. Depending on the tree species, leaf water content declined (red maple) and carbon‐based phenolics increased (sugar maple) on plants grown in an enriched CO₂ atmosphere. The only observed effect of elevated temperature on leaf phytochemistry was a reduction in leaf water content of sugar maple leaves. Gypsy moth larval responses were dependent on tree species. Larvae feeding on elevated CO₂‐grown red maple leaves had reduced growth, while temperature had no effect on the growth or consumption of larvae. No significant effects of either temperature or CO₂ concentration were observed for larvae feeding on sugar maple leaves. Our data demonstrate strong effects of CO₂ enrichment on leaf phytochemical constituents important to folivorous insects, while an elevated temperature largely has little effect. We conclude that alterations in leaf chemistry due to an elevated CO₂ atmosphere are more important in this plant–folivorous insect system than either the direct short‐term effects of temperature on insect performance or its indirect effects on leaf chemistry.     

Elevated CO2 reduces the nitrogen concentration of plant tissues

We summarize the impacts of elevated CO₂ on the N concentration of plant tissues and present data to support the hypothesis that reductions in the quality of plant tissue commonly occur when plants are grown under elevated CO₂. Synthesis of existing data showed an average 14% reduction of N concentrations in plant tissue generated under elevated CO₂ regimes. However, elevated CO₂ appeared to have different effects on the N concentrations of different plant types, as the reported reductions in N have been larger in C3 plants than in C4 plants and N₂-fixers. Under elevated CO₂ plants changed their allocation of N between above- and below-ground components: root N concentrations were reduced by an average of 9% compared to a 14% average reduction for above-ground tissues. Although the concentration of CO₂ treatments represented a significant source of variance for plant N concentration, no consistent trends were observed between them.     

Impact of elevated ozone concentration on growth,physiology, and yield of wheat (Triticum aestivum L.): a meta‐analysis

We quantitatively evaluated the effects of elevated concentration of ozone (O₃) on growth, leaf chemistry, gas exchange, grain yield, and grain quality relative to carbon‐filtered air (CF) by means of meta‐analysis of published data. Our database consisted of 53 peer‐reviewed studies published between 1980 and 2007, taking into account wheat type, O₃ fumigation method, rooting environment, O₃ concentration ([O₃]), developmental stage, and additional treatments such as drought and elevated carbon dioxide concentration ([CO₂]). The results suggested that elevated [O₃] decreased wheat grain yield by 29% (CI: 24–34%) and aboveground biomass by 18% (CI: 13–24%), where CI is the 95% confidence interval. Even in studies where the [O₃] range was between 31 and 59 ppb (average 43 ppb), there was a significant decrease in the grain yield (18%) and biomass (16%) relative to CF. Despite the increase in the grain protein content (6.8%), elevated [O₃] significantly decreased the grain protein yield (?18%). Relative to CF, elevated [O₃] significantly decreased photosynthetic rates (?20%), Rubisco activity (?19%), stomatal conductance (?22%), and chlorophyll content (?40%). For the whole plant, rising [O₃] induced a larger decrease in belowground (?27%) biomass than in aboveground (?18%) biomass. There was no significant response difference between spring wheat and winter wheat. Wheat grown in the field showed larger decreases in leaf photosynthesis parameters than wheat grown in < 5 L pots. Open‐top chamber fumigation induced a larger reduction than indoor growth chambers, when plants were exposed to elevated [O₃]. The detrimental effect was progressively greater as the average daily [O₃] increased, with very few exceptions. The impact of O₃ increased with developmental stages, with the largest detrimental impact during grain filling. Both drought and elevated [CO₂] significantly ameliorated the detrimental effects of elevated [O₃], which could be explained by a significant decrease in O₃ uptake resulting from decreased stomatal conductance.     

Effects of elevated O3, alone and in combination with elevated CO2, on tree leaf chemistry and insect herbivore performance: a meta-analysis

We reviewed the effects of elevated ozone (O₃), alone and in combination with elevated carbon dioxide (CO₂) on primary and secondary metabolites of trees and performance of insect herbivores by means of meta‐analysis. Our database consisted of 63 studies conducted on 22 species of trees and published between 1990 and 2005. Ozone alone had no overall effect on concentrations of carbohydrates or nutrients, whereas in combination with CO₂, elevated O₃ reduced nutrient concentrations and increased carbohydrate concentrations. In contrast to primary metabolites, concentrations of phenolics and terpenes were significantly increased by 16% and 8%, respectively, in response to elevated O₃. Effects of ozone in combination with elevated CO₂ were weaker than those of ozone alone on phenolics, but stronger than those of ozone alone on terpenes. The magnitude of secondary metabolite responses depended on the type of ozone exposure facility and increased in the following order: indoor growth chamber 3 than gymnosperms, as shifts in concentrations of carbohydrate and phenolics were observed in the former, but not in the latter. Elevated O₃ had positive effects on some indices of insect performance: pupal mass increased and larval development time shortened, but these effects were counteracted by elevated CO₂. Therefore, despite the observed increase in secondary metabolites, elevated O₃ tends to increase tree foliage quality for herbivores, but elevated CO₂ may alleviate these effects. Our meta‐analysis clearly demonstrated that effects of elevated O₃ alone on leaf chemistry and some indices of insect performance differed from those of O₃+CO₂, and therefore, it is important to study effects of several factors of global climate change simultaneously.     

Responses of cotton and wheat photosynthesis and growth to cyclic variation in carbon dioxide concentration

The carbon dioxide concentration in free air carbon dioxide enrichment (FACE) systems typically has rapid fluctuations. In our FACE system, power spectral analysis of CO₂ concentration measured every second with an open path analyzer indicated peaks in variation with a period of about one minute. I used open-top chambers to expose cotton and wheat plants to either a constant elevated CO₂ concentration of 180 ??mol mol^?1 above that of outside ambient air, or to the same mean CO₂ concentration, but with the CO₂ enrichment cycling between about 30 and 330 ??mol mol^?1 above the concentration of outside ambient air, with a period of one minute. Three short-term replicate plantings of cotton were grown in Beltsville, Maryland with these CO₂ concentration treatments imposed for 27-day periods over two summers, and one winter wheat crop was grown from sowing to maturity. In cotton, leaf gas-exchange measurements of the continuously elevated treatment and the fluctuating treatment indicated that the fluctuating CO₂ concentration treatment consistently resulted in substantial down-regulation of net photosynthetic rate (P _N) and stomatal conductance (g _s). Total shoot biomass of the vegetative cotton plants in the fluctuating CO₂ concentration treatment averaged 30% less than in the constantly elevated CO₂ concentration treatment at 27 days after planting. In winter wheat, leaf gas-exchange measurements also indicated that down-regulation of P _N and g _s occurred in flag leaves in the fluctuating CO₂ concentration treatment, but the effect was not as consistent in other leaves, nor as severe as found in cotton. However, wheat grain yields were 12% less in the fluctuating CO₂ concentration treatment compared with the constant elevated CO₂ concentration treatment. Comparison with wheat yields in chambers without CO₂ addition indicated a nonsignificant increase of 5% for the fluctuating elevated CO₂ concentration treatment, and a significant increase of 19% for the constant elevated treatment. The results suggest that treatments with fluctuating elevated CO₂ concentrations could underestimate plant growth at projected future atmospheric CO₂ concentrations.     

Elevated atmospheric CO2 effects on biomass production and soil carbon in conventional and conservation cropping systems

Increasing atmospheric CO₂ concentration has led to concerns about potential effects on production agriculture as well as agriculture's role in sequestering C. In the fall of 1997, a study was initiated to compare the response of two crop management systems (conventional and conservation) to elevated CO₂. The study used a split‐plot design replicated three times with two management systems as main plots and two CO₂ levels (ambient=375 μL L^?1 and elevated CO₂=683 μL L^?1) as split‐plots using open‐top chambers on a Decatur silt loam (clayey, kaolinitic, thermic Rhodic Paleudults). The conventional system was a grain sorghum (Sorghum bicolor (L.) Moench.) and soybean (Glycine max (L.) Merr.) rotation with winter fallow and spring tillage practices. In the conservation system, sorghum and soybean were rotated and three cover crops were used (crimson clover (Trifolium incarnatum L.), sunn hemp (Crotalaria juncea L.), and wheat (Triticum aestivum L.)) under no‐tillage practices. The effect of management on soil C and biomass responses over two cropping cycles (4 years) were evaluated. In the conservation system, cover crop residue (clover, sunn hemp, and wheat) was increased by elevated CO₂, but CO₂ effects on weed residue were variable in the conventional system. Elevated CO₂ had a greater effect on increasing soybean residue as compared with sorghum, and grain yield increases were greater for soybean followed by wheat and sorghum. Differences in sorghum and soybean residue production within the different management systems were small and variable. Cumulative residue inputs were increased by elevated CO₂ and conservation management. Greater inputs resulted in a substantial increase in soil C concentration at the 0–5 cm depth increment in the conservation system under CO₂‐enriched conditions. Smaller shifts in soil C were noted at greater depths (5–10 and 15–30 cm) because of management or CO₂ level. Results suggest that with conservation management in an elevated CO₂ environment, greater residue amounts could increase soil C storage as well as increase ground cover.     

Seasonal changes in the response of winter wheat to elevated atmospheric CO2 concentration grown in Open-Top Chambers and field tracking enclosures

Winter wheat was grown at ambient and elevated (ambient plus 350 μL L^–1) CO₂ concentrations in open top chambers and in field-tracking sun-lit climatized enclosures (elevated is 718 μL L^–1). There was no significant effect of CO₂ concentration on sheath, leaf and root biomass and leaf area in the early spring (January to April). 24-h canopy CO₂ exchange rate (CCER) was not significantly affected either. However, elevated CO₂ concentration increased CCER at midday, decreased evapotranspiration rate and increased instantaneous water-use-efficiency during early spring. Leaf, sheath and root nitrogen concentration per unit dry weight decreased and nonstructural carbohydrate concentration increased under elevated CO₂, and N-uptake per unit ground area decreased significantly (– 22%) towards the end of this period. These results contrast with results from the final harvest, when grain yield and biomass were increased by 19% under elevated CO₂. N concentration per dry weight was reduced by 5%, but N-uptake per unit ground area was significantly higher (+ 11%) for the elevated CO₂ treatment. 24-h and midday-CCER increased significantly more in late spring (period of 21 April to 30 May) (respectively by + 40% and 53%) than in the early spring (respectively 5% and 19%) in response to elevated CO₂. Midday evapotranspiration rate was reduced less by elevated CO₂ in the late spring (– 13%) than in early spring (– 21%). The CO₂ response of midday and 24-h CCER decreased again (+ 27% and + 23% resp.) towards the end of the growing season. We conclude that the low response to CO₂ concentration during the early spring was associated with a growth-restriction, caused by low temperature and irradiance levels. The reduction of nitrogen concentration, the increase of nonstructural carbohydrate, and the lower evapotranspiration indicated that CO₂ did have an effect towards the end of early spring, but not on biomass accumulation. Regression analysis showed that both irradiance and temperature affected the response to CO₂.     

Nitrogen application is required to realize wheat yield stimulation by elevated CO2 but will not remove the CO2‐induced reduction in grain protein concentration

Elevated CO₂ (eCO₂) generally promotes increased grain yield (GY) and decreased grain protein concentration (GPC), but the extent to which these effects depend on the magnitude of fertilization remains unclear. We collected data on the eCO₂ responses of GY, GPC and grain protein yield and their relationships with nitrogen (N) application rates across experimental data covering 11 field grown wheat (Triticum aestivum) cultivars studied in eight countries on four continents. The eCO₂‐induced stimulation of GY increased with N application rates up to ~200 kg/ha. At higher N application, stimulation of GY by eCO₂ stagnated or even declined. This was valid both when the yield stimulation was expressed as the total effect and using per ppm CO₂ scaling. GPC was decreased by on average 7% under eCO₂ and the magnitude of this effect did not depend on N application rate. The net effect of responses on GY and protein concentration was that eCO₂ typically increased and decreased grain protein yield at N application rates below and above ~100 kg/ha respectively. We conclude that a negative effect on wheat GPC seems inevitable under eCO₂ and that substantial N application rates may be required to sustain wheat protein yields in a world with rising CO₂.     

Effects of increased atmospheric CO2 on nutritional contents in poplar (Populus pseudo-simonii [Kitag.]) tissues and larval growth of gypsy moth (Lymantria dispar)

Changes in the concentrations of phytochemical compounds usually occur when plants are grown under elevated atmospheric CO₂. CO₂-induced changes in foliar chemistry tend to reduce leaf quality and may further affect insect herbivores. Increased atmospheric CO₂ also has a potential influence on decomposition because it causes variations in chemical components of plant tissues. To investigate the effects of increased atmospheric CO₂ on the nutritional contents of tree tissues and the activities of leaf-chewing forest insects, samples of Populus pseudo-simonii [Kitag.] grown in open-top chambers under ambient and elevated CO₂ (650 μmol mol^-1) conditions were collected for measuring concentrations of carbon, nitrogen, C : N ratio, soluble sugar and starch in leaves, barks, coarse roots (>2 mm in diameter) and fine roots (<2 mm in diameter). Gypsy moth (Lymantria dispar) larvae were reared on a single branch of experimental trees in a nylon bag with 1 mm 1 mm grid. The response of larval growth was observed in situ. Elevated CO₂ resulted in significant reduction in nitrogen concentration and increase in C : N ratio of all poplar tissues. In all tissues, total carbon contents were not affected by CO₂ treatments. Soluble sugar and nonstructural carbohydrate (TNC) in the poplar leaves significantly increased with CO₂ enrichment, whereas starch concentration increased only on partial sampling dates. Carbohydrate concentration in roots and barks was generally not affected by elevated CO₂, whereas soluble sugar contents in fine roots decreased in response to elevated CO₂. When second instar gypsy moth larvae consuming poplars grew under elevated CO₂ for the first 13 days, their body weight was 30.95% lower than that of larvae grown at ambient CO₂, but no significant difference was found when larvae were fed in the same treatment for the next 11 days. Elevated atmospheric CO₂ had adverse effects on the nutritional quality of Populus pseudo-simonii [Kitag.] tissues and the resultant variations in foliar chemical components had a significant but negative effect on the growth of early instar gypsy moth larvae.