Insects and fungi on a C3 sedge and a C4 grass exposed to elevated atmospheric CO2 concentrations in open-top chambers in the field

The effects of elevated atmospheric CO₂ concentration on plant-fungi and plant-insect interactions were studied in an emergent marsh in the Chesapeake Bay. Stands of the C₃ sedge Scirpus olneyi Grey, and the C₄ grass Spartina patens (Ait.) Muhl. have been exposed to elevated atmospheric CO₂ concentrations during each growing season since 1987. In August 1991 the severities of fungal infections and insect infestations were quantified. Shoot nitrogen concentration ([N]) and water content (WC) were determined. In elevated concentrations of atmospheric CO₂, 32% fewer S. olneyi plants were infested by insects, and there was a 37% reduction in the severity of a pathogenic fungal infection, compared with plants grown in ambient CO₂ concentrations. S. olneyi also had reduced [N], which correlated positively with the severities of fungal infections and insect infestations. Conversely, S. patens had increased WC but unchanged [N] in elevated concentrations of atmospheric CO₂ and the severity of fungal infection increased. Elevated atmospheric CO₂ concentration increased or decreased the severity of fungal infection depending on at least two interacting factors, [N] and WC; but it did not change the number of plants that were infected with fungi. In contrast, the major results for insects were that the number of plants infected with insects decreased, and that the amount of tissue that each insect ate also decreased.     

Sagebrush carbon allocation patterns and grasshopper nutrition: the influence of CO2 enrichment and soil mineral limitation

Summary Artemisia tridentata seedlings were grown under carbon dioxide concentrations of 350 and 650 l l^–1 and two levels of soil nutrition. In the high nutrient treatment, increasing CO₂ led to a doubling of shoot mass, whereas nutrient limitation completely constrained the response to elevated CO₂. Root biomass was unaffected by any treatment. Plant root/shoot ratios declined under carbon dioxide enrichment but increased under low nutrient availability, thus the ratio was apparently controlled by changes in carbon allocation to shoot mass alone. Growth under CO₂ enrichment increased the starch concentrations of leaves grown under both nutrient regimes, while increased CO₂ and low nutrient availability acted in concert to reduce leaf nitrogen concentration and water content. Carbon dioxide enrichment and soil nutrient limitation both acted to increase the balance of leaf storage carbohydrate versus nitrogen (C/N). The two treatment effects were significantly interactive in that nutrient limitation slightly reduced the C/N balance among the high-CO₂ plants. Leaf volatile terpene concentration increased only in the nutrient limited plants and did not follow the overall increase in leaf C/N ratio. Grasshopper consumption was significantly greater on host leaves grown under CO₂ enrichment but was reduced on leaves grown under low nutrient availability. An overall negative relationship of consumption versus leaf volatile concentration suggests that terpenes may have been one of several important leaf characteristics limiting consumption of the low nutrient hosts. Digestibility of host leaves grown under the high CO₂ treatment was significantly increased and was related to high leaf starch content. Grasshopper growth efficiency (ECI) was significantly reduced by the nutrient limitation treatment but co-varied with leaf water content.     

Infection with the parasitic angiosperm Striga hermonthica influences the response of the C3 cereal Oryza sativa to elevated CO2

Upland rice (Oryza sativa L.) was grown at both ambient (350 μmol mol^?1) and elevated (700 μmol mol^?1) CO₂ in either the presence or absence of the root hemi‐parasitic angiosperm Striga hermonthica (Del) Benth. Elevated CO₂ alleviated the impact of the parasite on host growth: biomass of infected rice grown at ambient CO₂ was 35% that of uninfected, control plants, while at elevated CO₂, biomass of infected plants was 73% that of controls. This amelioration occurred despite the fact that O. sativa grown at elevated CO₂ supported both greater numbers and a higher biomass of parasites per host than plants grown at ambient CO₂. The impact of infection on host leaf area, leaf mass, root mass and reproductive tissue mass was significantly lower in plants grown at elevated as compared with ambient CO₂. There were significant CO₂ and Striga effects on photosynthetic metabolism and instantaneous water‐use efficiency of O. sativa. The response of photosynthesis to internal [CO₂] (A/C_i curves) indicated that, at 45 days after sowing (DAS), prior to emergence of the parasites, uninfected plants grown at elevated CO₂ had significantly lower CO₂ saturated rates of photosynthesis, carboxylation efficiencies and ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco; EC 4.1.1.39) contents than uninfected, ambient CO₂‐grown O. sativa. In contrast, infection with S. hermonthica prevented down‐regulation of photosynthesis in O. sativa grown at elevated CO₂, but had no impact on photosynthesis of hosts grown at ambient CO₂. At 76 DAS (after parasites had emerged), however, infected plants grown at both elevated and ambient CO₂ had lower carboxylation efficiencies and Rubisco contents than uninfected O. sativa grown at ambient CO₂. The reductions in carboxylation efficiency (and Rubisco content) were accompanied by similar reductions in nitrogen concentration of O. sativa leaves, both before and after parasite emergence. There were no significant CO₂ or infection effects on the concentrations of soluble sugars in leaves of O. sativa, but starch concentration was significantly lower in infected plants at both CO₂ concentrations. These results demonstrate that elevated CO₂ concentrations can alleviate the impact of infection with Striga on the growth of C₃ hosts such as rice and also that infection can delay the onset of photosynthetic down‐regulation in rice grown at elevated CO₂.     

Comparison of photosynthetic acclimation to elevated CO2 and limited nitrogen supply in soybean

Plants grown at elevated CO₂ often acclimate such that their photosynthetic capacities are reduced relative to ambient CO₂-grown plants. Reductions in synthesis of photosynthetic enzymes could result either from reduced photosynthetic gene expression or from reduced availability of nitrogen-containing substrates for enzyme synthesis. Increased carbohydrate concentrations resulting from increased photosynthetic carbon fixation at elevated CO₂ concentrations have been suggested to reduce the expression of photosynthetic genes. However, recent studies have also suggested that nitrogen uptake may be depressed by elevated CO₂, or at least that it is not increased enough to keep pace with increased carbohydrate production. This response could induce a nitrogen limitation in elevated-CO₂ plants that might account for the reduction in photosynthetic enzyme synthesis. If CO₂ acclimation were a response to limited nitrogen uptake, the effects of elevated CO₂ and limiting nitrogen supply on photosynthesis and nitrogen allocation should be similar. To test this hypothesis we grew non-nodulating soybeans at two levels each of nitrogen and CO₂ concentration and measured leaf nitrogen contents, photosynthetic capacities and Rubisco contents. Both low nitrogen and elevated CO₂ reduced nitrogen as a percentage of total leaf dry mass but only low nitrogen supply produced significant decreases in nitrogen as a percentage of leaf structural dry mass. The primary effect of elevated CO₂ was to increase non-structural carbohydrate storage rather than to decrease nitrogen content. Both low nitrogen supply and elevated CO₂ also decreased carboxylation capacity (V_cmax) and Rubisco content per unit leaf area. However, when V_cmax and Rubisco content were expressed per unit nitrogen, low nitrogen supply generally caused them to increase whereas elevated CO₂ generally caused them to decrease. Finally, elevated CO₂ significantly increased the ratio of RuBP regeneration capacity to V_cmax whereas neither nitrogen supply nor plant age had a significant effect on this parameter. We conclude that reductions in photosynthetic enzyme synthesis in elevated CO₂ appear not to result from limited nitrogen supply but instead may result from feedback inhibition by increased carbohydrate contents.     

Elevated atmospheric carbon dioxide and ozone alter soybean diseases at SoyFACE

Human driven changes in the Earth's atmospheric composition are likely to alter plant disease in the future. We evaluated the effects of elevated carbon dioxide (CO₂) and ozone (O₃) on three economically important soybean diseases (downy mildew, Septoria brown spot and sudden death syndrome‐SDS) under natural field conditions at the soybean free air concentration enrichment (SoyFACE) facility. Disease incidence and/or severity were quantified from 2005 to 2007 using visual surveys and digital image analysis, and changes were related to microclimatic variability and to structural and chemical changes in soybean host plants. Changes in atmospheric composition altered disease expression, but responses of the three pathosystems varied considerably. Elevated CO₂ alone or in combination with O₃ significantly reduced downy mildew disease severity (measured as area under the disease progress curve‐AUDPC) by 39–66% across the 3 years of the study. In contrast, elevated CO₂ alone or in combination with O₃ significantly increased brown spot severity in all 3 years, but the increase was small in magnitude. When brown spot severity was assessed in relation to differences in canopy height induced by the atmospheric treatments, disease severity increased under combined elevated CO₂ and O₃ treatment in only one of the 3 years. The atmospheric treatments had no effect on the incidence of SDS or brown spot throughout the study. Higher precipitation during the 2006 growing season was associated with increased AUDPC severity across all treatments by 2.7 and 1.4 times for downy mildew and brown spot, respectively, compared with drought conditions in 2005. In the 2 years with similar precipitation, the higher daily temperatures in the late spring of 2007 were associated with increased severity of downy mildew and brown spot. Elevated CO₂ and O₃ induced changes in the soybean canopy density and leaf age likely contributed to the disease expression modifications.     

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.     

Growth, senescence and water use efficiency of spring oilseed rape (Brassica napus L. cv. Mozart) grown in a factorial combination of nitrogen supply and elevated CO2

Atmospheric CO₂ enrichment is expected to affect the resource use efficiency of C3 plants with respect to water, nutrients and light in an interactive manner. The responses of oilseed rape (OSR) to elevated CO₂ have not much been addressed. Since the crop has low nitrogen use efficiency, the interactive effects of CO₂ enrichment and nitrogen supply deserve particular attention.Spring OSR was grown in climate chambers simulating the seasonal increments of day length and temperature in South-Western Germany. Three levels of N fertilisation representing 75, 150 and 225 kg ha⁻¹ and two CO₂ concentrations (380 and 550 μmol mol⁻¹) were used to investigate changes in source-sink relationships, plant development and senescence, water use efficiency of the dry matter production (WUE_prod.), allocation patterns to different fractions, growth, yield and seed oil contents. Seven harvests were performed between 72 and 142 days after sowing (DAS).Overall, plant performance in the chambers was comparable to the development under field conditions. While CO₂ responses were small in the plants receiving lowest N-levels, several significant N × CO₂ interactions were observed in the other treatments. Increasing the N availability resulted in longer flowering windows, which were furthermore extended at elevated CO₂ concentrations. Nevertheless, significantly less biomass was allocated to reproductive structures under elevated CO₂, while the vegetative C-storing organs continued to grow. At the final harvest shoot mass of the CO₂ exposed plants had increased by 9, 8 and 15% in the low, medium and high N treatments. Root growth was increased even more by 17, 43 and 33%, respectively and WUE_prod. increased by 23, 42 and 35%. At the same time, seed oil contents were significantly reduced by CO₂ enrichment in the treatments with ample N supply.Obviously, under high N-supply, the CO₂ fertilisation induced exaggerated growth of vegetative tissues at the expense of reproductive structures. The interruption of source-sink relationships stimulated the formation of side shoots and flowers (branching out). While direct effects of elevated CO₂ on flowering can be excluded, we assume that the increased growth under high N and CO₂ supply created nutrient imbalances which hence affected flowering and seed set.Nevertheless, the final seed macronutrient concentrations were slightly increased by elevated CO₂, indicating that remobilisation of nutrients from the sources (leaves) to the sinks (seeds) remained effective. These findings were supported by the lower nitrogen concentrations in senescing leaves and probably increased N remobilisation to other plant parts under elevated concentrations of CO₂. All the same, CO₂ enrichment caused a decline in seed oil contents, which may translate into a reduced crop quality.     

Short-term decomposition of litter produced by plants grown in ambient and elevated atmospheric CO2 concentrations

The effects of elevated atmospheric CO₂ (ambient + 340 μmol mol^–1) on above-ground litter decomposition were investigated over a 6-week period using a field-based mesocosm system. Soil respiratory activity in mesocosms incubated in ambient and elevated atmospheric CO₂ concentrations were not significantly different (t-test, P > 0.05) indicating that there were no direct effects of elevated atmospheric CO₂ on litter decomposition. A study of the indirect effects of CO₂ on soil respiration showed that soil mesocosms to which naturally senescent plant litter had been added (0.5% w/w) from the C₃ sedge Scirpus olneyi grown in elevated atmospheric CO₂ was reduced by an average of 17% throughout the study when compared to soil mesocosms to which litter from Scirpus olneyi grown in ambient conditions had been added. In contrast, similar experiments using senescent material from the C₄ grass Spartina patens showed no difference in soil respiration rates between mesocosms to which litter from plants grown in elevated or ambient CO₂ conditions had been added. Analysis of the C:N ratio and lignin content of the senescent material showed that, while the C:N ratio and lignin content of the Spartina patens litter did not vary with atmospheric CO₂ conditions, the C:N ratio (but not the lignin content) of the litter from Scirpus olneyi was significantly greater (t-test;P < 0.05) when derived from plants grown under elevated CO₂ (105:1 compared to 86:1 for litter derived from Scirpus olneyi grown under ambient conditions). The results suggest that the increased C:N ratio of the litter from the C₃ plant Scirpus olneyi grown under elevated CO₂ led to the lower rates of biodegradation observed as reduced soil respiration in the mesocosms. Further long-term experiments are now required to determine the effects of elevated CO₂ on C partitioning in terrestrial ecosystems.     

Aphid suitability for the predatory hoverfly Episyrphus balteatus altered with elevating atmospheric CO2 and sinigrin

The fitness of natural enemies should be altered in response to changes in herbivore quality induced by the impact of increased atmospheric CO₂ levels on plants. We studied the effect of different CO₂ levels on the aphid predator Episyrphus balteatus DeGeer fed either specialist or generalist aphids reared on either of two host plants under laboratory conditions. In the host plant that contains sinigrin (black mustard), elevated CO₂ increased the sinigrin content of both host plant and the specialist aphid, but reduced the already very low levels in the generalist aphid. Predator development time increased with elevated CO₂, while fecundity decreased. Consequently, individual fitness decreased slightly with increasing atmospheric CO₂. Sinigrin significantly decreased fecundity and increased development time of the predator. As a result, fitness was significantly lower too. The consumption rate was influenced significantly by plant and prey solely and the interactions of host plant × prey type and CO₂ level × prey type. Further research on the effects of climate change parameters (e.g. greenhouse gases such as CO₂, ozone (O₃) and nitrogen dioxide (NO₂), etc.) separately and jointly under controlled environmental conditions will help to understand the nature and direction of their effects on natural enemies as part of the tritrophic system.     

Assimilation,respiration and allocation of carbon inPlantago major as affected by atmospheric CO2 levels

The response ofPlantago major ssp,pleiosperma plants, grown on nutrient solution in a climate chamber, to a doubling of the ambient atmospheric CO₂ concentration was investigated. Total dry matter production was increased by 30% after 3 weeks of exposure, due to a transient stimulation of the relative growth rate (RGR) during the first 10 days. Thereafter RGR returned to the level of control plants. Photosynthesis, expressed per unit leaf area, was stimulated during the first two weeks of the experiment, thereafter it dropped and nearly reached the level of the control plants. Root respiration was not affected by increased atmospheric CO₂ levels, whereas shoot, dark respiration was stimulated throughout the experimental period. Dry matter allocation over leaves stems and roots was not affected by the CO₂ level. SLA was reduced by 10%, which can partly be explained by an increased dry matter content of the leaves. Both in the early and later stages of the experiment, shoot respiration accounted for a larger part of the carbon budget in plants grown at elevated atmospheric CO₂. Shifts in the total carbon budget were mainly due to the effects on shoot respiration. Leaf growth accounted for nearly 50% of the C budget at all stages of the experiment and in both treatments.Abbreviations LAR leaf area ratio - LWR leaf weight ratio - RGR relative growth rate - R/S root to shoot ratio - RWR root weight ratio - SLA specific leaf area - SWR stem weight ratio     

Controls of biomass partitioning between roots and shoots: Atmospheric CO2 enrichment and the acquisition and allocation of carbon and nitrogen in wild radish

Summary The effects of CO₂ enrichment on plant growth, carbon and nitrogen acquisition and resource allocation were investigated in order to examine several hypotheses about the mechanisms that govern dry matter partitioning between shoots and roots. Wild radish plants (Raphanus sativus × raphanistrum) were grown for 25 d under three different atmospheric CO₂ concentrations (200 ppm, 330 ppm and 600 ppm) with a stable hydroponic 150 mol 1^–1 nitrate supply. Radish biomass accumulation, photosynthetic rate, water use efficiency, nitrogen per unit leaf area, and starch and soluble sugar levels in leaves increased with increasing atmospheric CO₂ concentration, whereas specific leaf area and nitrogen concentration of leaves significantly decreased. Despite substantial changes in radish growth, resource acquisition and resource partitioning, the rate at which leaves accumulated starch over the course of the light period and the partitioning of biomass between roots and shoots were not affected by CO₂ treatment. This phenomenon was consistent with the hypothesis that root/shoot partitioning is related to the daily rate of starch accumulation by leaves during the photoperiod, but is inconsistent with hypotheses suggesting that root/shoot partitioning is controlled by some aspect of plant C/N balance.     

Elevated atmospheric CO2 and increased nitrogen deposition: effects on C and N metabolism and growth of the peat moss Sphagnum recurvum P. Beauv. var. mucronatum (Russ.) Warnst

Sphagnum bogs play an important role when considering the impacts of global change on global carbon and nitrogen cycles. Sphagnum recurvum P. Beauv. var. mucronatum (Russ.) was grown at 360 (ambient) and 700 μL L^?1 (elevated) atmospheric [CO₂] in combination with different nitrogen deposition rates (6, 15, 23 g N m^?2 y^?1), in a short‐ and long‐term growth chamber experiment. After 6 months, elevated atmospheric [CO₂] in combination with the lowest nitrogen deposition rate, increased plant dry mass by 17%. In combination with a high nitrogen deposition rate, biomass production was not significantly stimulated. At the start of the experiment, photosynthesis was stimulated by elevated atmospheric [CO₂], but it was downregulated to control levels after three days of exposure. Elevated [CO₂] substantially reduced dark respiration, which resulted in a continuous increase in soluble sugar content in capitula. Differences in growth response among different nitrogen and CO₂ treatments could not be related to measured carbon exchange rates, which was mainly due to interference of microbial respiration. Doubling atmospheric [CO₂] reduced total nitrogen content in capitula but not in stems at all nitrogen deposition rates. Reduction in total nitrogen content coincided with a decrease in amino acids, but soluble protein levels remained unaffected. Thus, elevated [CO₂] induced a substantial shift in the partitioning of nitrogen compounds in capitula. Soluble sugar concentration was negatively correlated with total nitrogen content, which implies that the reduction in amino acid content in capitula, exposed to elevated [CO₂], might be caused by the accumulation of soluble sugars. Growth was not stimulated by increased nitrogen deposition. High nitrogen deposition, resulting in a capitulum nitrogen content in excess of 15 mg g^?1 dw, was detrimental to photosynthesis, reduced water content and induced necrosis. We propose a capitulum nitrogen content of 15 mg g^?1 dw as a possible bioindicator for the detection of nitrogen pollution stress in oligo‐mesotrophic peat bog ecosystems. At the lowest nitrogen deposition level, nitrogen recovery was higher than 100%, which indicates substantial dry deposition and/or gaseous nitrogen fixation by bacteria, associated with Sphagnum. Increasing nitrogen deposition rates decreased nitrogen recovery percentages, which indicates reduced efficiency of nitrogen fixation.     

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 elevated CO2 and transgenic Bt cotton on plant chemistry, performance, and feeding of an insect herbivore, the cotton bollworm

Effects of elevated atmospheric CO₂ (double‐ambient CO₂) on the growth and metabolism of cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), fed on transgenic Bacillus thuringiensis (Berliner) (Bt) cotton [Cry1A(c)], grown in open‐top chambers, were studied. Two levels of CO₂ (ambient and double‐ambient) and two cotton cultivars (non‐transgenic Simian‐3 and transgenic GK‐12) were deployed in a completely randomized design with four treatment combinations, and the cotton bollworm was reared on each treatment simultaneously. Plants of both cotton cultivars had lower nitrogen and higher total non‐structural carbohydrates (TNC), TNC:Nitrogen ratio, condensed tannin, and gossypol under elevated CO₂. Elevated CO₂ further resulted in a significant decrease in Bt toxin level in GK‐12. The changes in chemical components in the host plants due to increased CO₂ significantly affected the growth parameters of H. armigera. Both transgenic Bt cotton and elevated CO₂ resulted in a reduced body mass, lower fecundity, decreased relative growth rate (RGR), and decreased mean relative growth rate in the bollworms. Larval life‐span was significantly longer for H. armigera fed transgenic Bt cotton. Significantly reduced larval, pupal, and adult moth weights were observed in the bollworms fed elevated CO₂‐grown transgenic Bt cotton compared with those of bollworms reared on non‐transgenic cotton, regardless of the CO₂ level. The efficiency of conversion of ingested food and of digested food of the bollworm were significantly reduced when fed transgenic Bt cotton, but there was no significant CO₂ or CO₂× cotton cultivar interaction. Approximate digestibility of larvae reared on transgenic cotton grown in elevated CO₂ was higher compared to that of larvae fed non‐transgenic cotton grown at ambient CO₂. The damage inflicted by cotton bollworm on cotton, regardless of the presence or absence of insecticidal genes, is predicted to be more serious under elevated CO₂ conditions because of individual compensatory feeding on host plants caused by nitrogen deficiency.     

Plant nitrogen acquisition and interactions under elevated carbon dioxide: impact of endophytes and mycorrhizae

Both endophytic and mycorrhizal fungi interact with plants to form symbiosis in which the fungal partners rely on, and sometimes compete for, carbon (C) sources from their hosts. Changes in photosynthesis in host plants caused by atmospheric carbon dioxide (CO₂) enrichment may, therefore, influence those mutualistic interactions, potentially modifying plant nutrient acquisition and interactions with other coexisting plant species. However, few studies have so far examined the interactive controls of endophytes and mycorrhizae over plant responses to atmospheric CO₂ enrichment. Using Festuca arundinacea Schreb and Plantago lanceolata L. as model plants, we examined the effects of elevated CO₂ on mycorrhizae and endophyte (Neotyphodium coenophialum) and plant nitrogen (N) acquisition in two microcosm experiments, and determined whether and how mycorrhizae and endophytes mediate interactions between their host plant species. Endophyte‐free and endophyte‐infected F. arundinacea varieties, P. lanceolata L., and their combination with or without mycorrhizal inocula were grown under ambient (400 μmol mol⁻¹) and elevated CO₂ (ambient + 330 μmol mol⁻¹). A ¹⁵N isotope tracer was used to quantify the mycorrhiza‐mediated plant acquisition of N from soil. Elevated CO₂ stimulated the growth of P. lanceolata greater than F. arundinacea, increasing the shoot biomass ratio of P. lanceolata to F. arundinacea in all the mixtures. Elevated CO₂ also increased mycorrhizal root colonization of P. lanceolata, but had no impact on that of F. arundinacea. Mycorrhizae increased the shoot biomass ratio of P. lanceolata to F. arundinacea under elevated CO₂. In the absence of endophytes, both elevated CO₂ and mycorrhizae enhanced ¹⁵N and total N uptake of P. lanceolata but had either no or even negative effects on N acquisition of F. arundinacea, altering N distribution between these two species in the mixture. The presence of endophytes in F. arundinacea, however, reduced the CO₂ effect on N acquisition in P. lanceolata, although it did not affect growth responses of their host plants to elevated CO₂. These results suggest that mycorrhizal fungi and endophytes might interactively affect the responses of their host plants and their coexisting species to elevated 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.     

Sagebrush and grasshopper responses to atmospheric carbon dioxide concentration

Summary Seed- and clonally-propagated plants of Big Sagebrush (Artemisia tridentata var.tridentata) were grown under atmospheric carbon dioxide regimes of 270, 350 and 650 μl l⁻¹ and fed toMelanoplus differentialis andM. sanguinipes grasshoppers. Total shrub biomass significantly increased as carbon dioxide levels increased, as did the weight and area of individual leaves. Plants grown from seed collected in a single population exhibited a 3–5 fold variation in the concentration of leaf volatile mono- and sesquiterpenes, guaianolide sesquiterpene lactones, coumarins and flavones within each CO₂ treatment. The concentration of leaf allelochemicals did not differ significantly among CO₂ treatments for these seed-propagated plants. Further, when genotypic variation was controlled by vegetative propagation, allelochemical concentrations also did not differ among carbon dioxide treatments. On the other hand, overall leaf nitrogen concentration declined significantly with elevated CO₂. Carbon accumulation was seen to dilute leaf nitrogen as the balance of leaf carbon versus nitrogen progressively increased as CO₂ growth concentration increased. Grasshopper feeding was highest on sagebrush leaves grown under 270 and 650 μl l⁻¹ CO₂, but varied widely within treatments. Leaf nitrogen concentration was an important positive factor in grasshopper relative growth but had no overall effect on consumption. Potential compensatory consumption by these generalist grasshoppers was apparently limited by the sagebrush allelochemicals. Insects with a greater ability to feed on chemically defended host plants under carbon dioxide enrichment may ultimately consume leaves with a lower nitrogen concentration but the same concentration of allelochemicals. Compensatory feeding may potentially increase the amount of dietary allelochemicals ingested for each unit of nitrogen consumed.     

Plant responses to atmospheric carbon dioxide enrichment: interactions with some soil and atmospheric conditions

In general, C₃ plant species are more responsive to atmospheric carbon dioxide (CO₂) enrichment than C₄-plants. Increased relative growth rate at elevated CO₂ primarily relates to increased Net Assimilation Rate (NAR), and enhancement of net photosynthesis and reduced photorespiration. Transpiration and stomatal conductance decrease with elevated CO₂, water use efficiency and shoot water potential increase, particularly in plants grown at high soil salinity. Leaf area per plant and leaf area per leaf may increase in an early growth stage with increased CO₂, after a period of time Leaf Area Ratio (LAR) and Specific Leaf Area (SLA) generally decrease. Starch may accumulate with time in leaves grown at elevated CO₂. Plants grown under salt stress with increased (dark) respiration as a sink for photosynthates, may not show such acclimation to increased atmospheric CO₂ levels. Plant growth may be stimulated by atmospheric carbon dioxide enrichment and reduced by enhanced UV-B radiation but the limited data available on the effect of combined elevated CO₂ and ultraviolet B (280–320 nm) (UV-B) radiation allow no general conclusion. CO₂-induced increase of growth rate can be markedly modified at elevated UV-B radiation. Plant responses to elevated atmospheric CO₂ and other environmental factors such as soil salinity and UV-B tend to be species-specific, because plant species differ in sensitivity to salinity and UV-B radiation, as well as to other environmental stress factors (drought, nutrient deficiency). Therefore, the effects of joint elevated atmospheric CO₂ and increased soil salinity or elevated CO₂ and enhanced UV-B to plants are physiologically complex.     

Elevated CO2 alleviates the impact of drought on barley improving water status by lowering stomatal conductance and delaying its effects on photosynthesis

We analysed the impact of elevated CO₂ on water relations, water use efficiency and photosynthetic gas exchange in barley (Hordeum vulgare L.) under wet and drying soil conditions. Soil moisture was less depleted under elevated compared to ambient [CO₂]. Elevated CO₂ had no significant effect on the water relations of irrigated plants, except on whole plant hydraulic conductance, which was markedly decreased at elevated compared to ambient CO₂ concentrations. The values of relative water content, water potential and osmotic potential were higher under elevated CO₂ during the entire drought period. The better water status of water-limited plants grown at elevated CO₂ was the result of stomatal control rather than of osmotic adjustment. Despite the low stomatal conductance produced by elevated CO₂, net photosynthesis was higher under elevated than ambient CO₂ concentrations. With water shortage, photosynthesis was maintained for longer at higher rates under elevated CO₂. The reduction of stomatal conductance and therefore transpiration, and the enhancement of carbon assimilation by elevated CO₂, increased instantaneous and whole plant water use efficiency in both irrigated and droughted plants. Thus, the metabolism of barley plants grown under elevated CO₂ and moderate or mild water deficit conditions is benefited by increased photosynthesis and lower transpiration. The reduction in plant water use results in a marked increase in soil water content which delays the onset and severity of water deficit.     

Effects of elevated CO2 and temperature on development and consumption rates of Octotoma championi and O. scabripennis feeding on Lantana camara

We carried out a factorial experiment to explore the effect of doubled CO₂ concentration and a 3 °C temperature increase on the development of a complete generation of the beetles Octotoma championi Baly and O. scabripennis Guérin‐Méneville (Coleoptera: Chrysomelidae). These species are biological control agents of Lantana camara L. (Verbenaceae), with a leaf‐mining larval phase and free‐living, leaf‐chewing adults. Plants grown at elevated CO₂ had enhanced above‐ground biomass, thicker leaves, reduced nitrogen concentration, and increased C:N ratios. Under the high temperature treatment, plants grown at ambient CO₂ suffered wilting and premature leaf loss, despite daily watering; this effect was ameliorated at elevated CO₂. The wilting of plants in the ambient CO₂/high temperature treatment reduced the emergence success of the beetles, particularly O. championi. Development time was accelerated by approximately 10–13 days at the higher temperature, but was not affected by CO₂. Neither CO₂ nor temperature affected adult beetle weight. Consumption rates of free‐living beetles were not affected by either CO₂ or temperature. By contrast, in the short‐term trials using excised foliage, beetles given no choice between ambient and elevated CO₂‐grown foliage, consumed more from ambient plants. When beetles were offered a choice between foliage grown at the two CO₂ levels, O. championi did not display a significant preference but O. scabripennis consumed more ambient CO₂‐grown foliage when feeding at the lower temperature. This study indicates that under future conditions of higher temperatures, amelioration of water stress in host plants growing in elevated CO₂ may benefit some endophagous insects by reducing premature leaf loss. Under some circumstances, this benefit may outweigh the deleterious effects of lower leaf nitrogen. Our results also indicate that foliage consumption under elevated CO₂ by mobile, adult insects on whole plants may not be significantly increased, as was previously indicated by short‐term experiments using excised foliage.