Interactions of elevated carbon dioxide and temperature with aphid feeding on transgenic oilseed rape: Are Bacillus thuringiensis (Bt) plants more susceptible to nontarget herbivores in future climate?

Climate change factors such as elevated carbon dioxide (CO₂) and temperature typically affect carbon (C) and nitrogen (N) dynamics of crop plants and the performance of insect herbivores. Insect‐resistant transgenic plants invest some nutrients to the production of specific toxic proteins [i.e. endotoxins from Bacillus thuringiensis (Bt)], which could alter the C–N balance of these plants, especially under changed abiotic conditions. Aphids are nonsusceptible to Lepidoptera‐targeted Bt Cry1Ac toxin and they typically show response to abiotic conditions, and here we sought to discover whether they might perform differently on compositionally changed Bt oilseed rape. Bt oilseed rape had increased N content in the leaves coupled with reduced total C compared with its nontransgenic counterpart, but in general the C : N responses of both plant types to elevated CO₂ and temperature were similar. Elevated CO₂ decreased N content and increased C : N ratio of both plant types. Elevated temperature increased C and N contents, total chlorophyll and carotenoid concentrations under ambient CO₂, but decreased these under elevated CO₂. In addition, soluble sugars were increased and starch decreased by elevated temperature under ambient but not under elevated CO₂, whereas photosynthesis was decreased in plants grown under elevated temperature in both CO₂ levels. Myzus persicae, a generalist aphid species, responded directly to elevated temperature with reduced developmental time and decreased adult and progeny weights, whereas the development of the Brassica specialist Brevicoryne brassicae was less affected. Feeding by M. persicae resulted in an increase in the N content of oilseed rape leaves under ambient CO₂, indicating the potential of herbivore feeding itself to cause allocation changes. The aphids performed equally well on both plant types despite the differences between C–N ratios of Bt and non‐Bt oilseed rape, revealing the absence of plant composition‐related effects on these pests under elevated CO₂, elevated temperature or combined elevated CO₂ and temperature conditions.     

Effects of elevated CO2 on five plant-aphid interactions

We investigated interactions between five species of phloem-feeding aphids (Homoptera: Aphididae) and their host plants at elevated CO₂; Acyrthosiphon pisum (Harris) on Vicia faba L., Aphis nerii Boyer de Fonscolombe on Asclepias syriaca L., Aphis oenotherae Oestlund on Oenothera biennis L., Aulacorthum solani (Kaltenbach) on Nicotiana sylvestris Speg. & Comes and Myzus persicae (Sulzer) on Solanum dulcamara L. Host plants grown at elevated CO₂ generally had greater biomass, leaf area and C:N ratios than those grown at ambient CO₂, while plants with aphids had lower biomass and leaf area than those without aphids.The responses of aphid populations to elevated CO₂ were species-specific with one species increasing (M. persicae), one decreasing (A. pisum), and the other three being unaffected. CO₂ treatment did not affect the proportion of alate individuals produced. In general, aphid abundance was not significantly related to foliar nitrogen concentration.We performed separate analyses to test whether either aphid presence or aphid abundance modified the response of host plants to elevated CO₂. In terms of aphid presence, only three of the potential 15 interactions (five aphid species x three plant traits) were significant; A. solani slightly modified the response of the plant biomass to elevated CO₂ and M. persicae affected the response of leaf area and allocation. In terms of aphid abundance, only two of the potential 15 interactions were significant with A. nerii modifying the plant response to CO₂ in terms of total leaf area and allocation.We conclude that, in contrast to other insect groups such as leaf chewers, populations of most phloem-feeders may not be negatively affected by increased CO₂ concentrations in the future. The reasons for this difference include the possibility that aphids may be able to compensate for changes in host plant quality by altering feeding behaviour or by synthesizing amino acids. In addition, there is little evidence that aphid herbivory, even at high levels, will substantially modify the response of plants to elevated CO₂.     

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

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

N2 fixation by Acacia species increases under elevated atmospheric CO2

In the present study the effect of elevated CO₂ on growth and nitrogen fixation of seven Australian Acacia species was investigated. Two species from semi‐arid environments in central Australia (Acacia aneura and A. tetragonophylla) and five species from temperate south‐eastern Australia (Acacia irrorata, A. mearnsii, A. dealbata, A. implexa and A. melanoxylon) were grown for up to 148 d in controlled greenhouse conditions at either ambient (350 µmol mol^?1) or elevated (700 µmol mol^?1) CO₂ concentrations. After establishment of nodules, the plants were completely dependent on symbiotic nitrogen fixation. Six out of seven species had greater relative growth rates and lower whole plant nitrogen concentrations under elevated versus normal CO₂. Enhanced growth resulted in an increase in the amount of nitrogen fixed symbiotically for five of the species. In general, this was the consequence of lower whole‐plant nitrogen concentrations, which equate to a larger plant and greater nodule mass for a given amount of nitrogen. Since the average amount of nitrogen fixed per unit nodule mass was unaltered by atmospheric CO₂, more nitrogen could be fixed for a given amount of plant nitrogen. For three of the species, elevated CO₂ increased the rate of nitrogen fixation per unit nodule mass and time, but this was completely offset by a reduction in nodule mass per unit plant mass.     

Elevated [CO2] and increased N supply reduce leaf disease and related photosynthetic impacts on Solidago rigida

To evaluate whether leaf spot disease and related effects on photosynthesis are influenced by increased nitrogen (N) input and elevated atmospheric CO₂ concentration ([CO₂]), we examined disease incidence and photosynthetic rate of Solidago rigida grown in monoculture under ambient or elevated (560 μmol mol⁻¹) [CO₂] and ambient or elevated (+4 g N m⁻² year⁻¹) N conditions in a field experiment in Minnesota, USA. Disease incidence was lower in plots with either elevated [CO₂] or enriched N (−57 and −37%, respectively) than in plots with ambient conditions. Elevated [CO₂] had no significant effect on total plant biomass, or on photosynthetic rate, but reduced tissue%N by 13%. In contrast, N fertilization increased both biomass and total plant N by 70%, and as a consequence tissue%N was unaffected and photosynthetic rate was lower on N fertilized plants than on unfertilized plants. Regardless of treatment, photosynthetic rate was reduced on leaves with disease symptoms. On average across all treatments, asymptomatic leaf tissue on diseased leaves had 53% lower photosynthetic rate than non-diseased leaves, indicating that the negative effect from the disease extended beyond the visual lesion area. Our results show that, in this instance, indirect effects from elevated [CO₂], i.e., lower disease incidence, had a stronger effect on realized photosynthetic rate than the direct effect of higher [CO₂].     

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.     

Competition for nitrogen between Pinus sylvestris and ectomycorrhizal fungi generates potential for negative feedback under elevated CO2

We investigated fungal species-specific responses of ectomycorrhizal (ECM) Scots pine (Pinus sylvestris) seedlings on growth and nutrient acquisition together with mycelial development under ambient and elevated CO₂. Each seedling was associated with one of the following ECM species: Hebeloma cylindrosporum, Laccaria bicolor, Suillus bovinus, S. luteus, Piloderma croceum, Paxillus involutus, Boletus badius, or non-mycorrhizal, under ambient, and elevated CO₂ (350 or 700 μl l⁻¹ CO₂); each treatment contained six replicates. The trial lasted 156 days. During the final 28 days, the seedlings were labeled with ¹⁴CO₂. We measured hyphal length, plant biomass, ¹⁴C allocation, and plant nitrogen and phosphorus concentration. Almost all parameters were significantly affected by fungal species and/or CO₂. There were very few significant interactions. Elevated CO₂ decreased shoot-to-root ratio, most strongly so in species with the largest extraradical mycelium. Under elevated CO₂, ECM root growth increased significantly more than hyphal growth. Extraradical hyphal length was significantly negatively correlated with shoot biomass, shoot N content, and total plant N uptake. Root dry weight was significantly negatively correlated with root N and P concentration. Fungal sink strength for N strongly affected plant growth through N immobilization. Mycorrhizal fungal-induced progressive nitrogen limitation (PNL) has the potential to generate negative feedback with plant growth under elevated CO₂. Responsible Editor: Herbert Johannes Kronzucker     

Legume presence increases photosynthesis and N concentrations of co-occurring non-fixers but does not modulate their responsiveness to carbon dioxide enrichment

Legumes, with the ability to fix atmospheric nitrogen (N), may help alleviate the N limitations thought to constrain plant community response to elevated concentrations of atmospheric carbon dioxide (CO₂). To address this issue we assessed: (1) the effects of the presence of the perennial grassland N₂ fixer, Lupinus perennis, on biomass accumulation and plant N concentrations of nine-species plots of differing plant composition; (2) leaf-level physiology of co-occurring non-fixing species (Achillea millefolium, Agropyron repens, Koeleria cristata) in these assemblages with and without Lupinus; (3) the effects of elevated CO₂ on Lupinus growth and symbiotic N₂ fixation in both monoculture and the nine-species assemblages; and (4) whether assemblages containing Lupinus exhibit larger physiological and growth responses to elevated CO₂ than those without. This study was part of a long-term grassland field experiment (BioCON) that controls atmospheric CO₂ at current ambient and elevated (560 µmol mol^–1) concentrations using free-air CO₂ enrichment. Nine-species plots with Lupinus had 32% higher whole plot plant N concentrations and 26% higher total plant N pools than those without Lupinus, based on both above and belowground measurements. Co-occurring non-fixer leaf N concentrations increased 22% and mass-based net photosynthetic rates increased 41% in plots containing Lupinus compared to those without. With CO₂ enrichment, Lupinus monocultures accumulated 32% more biomass and increased the proportion of N derived from fixation from 44% to 57%. In nine-species assemblages, Lupinus N derived from fixation increased similarly from 43% to 54%. Although Lupinus presence enhanced photosynthetic rates and leaf N concentrations of co-occurring non-fixers, and increased overall plant N pools, Lupinus presence did not facilitate stronger photosynthetic responses of non-fixing species or larger growth responses of overall plant communities to elevated CO₂. Non-fixer leaf N concentrations declined similarly in response to elevated CO₂ with and without Lupinus present and the relationship between net photosynthesis and leaf N was not affected by Lupinus presence. Regardless of the presence or absence of Lupinus, CO₂ enrichment resulted in reduced leaf N concentrations and rates of net photosynthesis.     

Increased allocation to external hyphae of arbuscular mycorrhizal fungi under CO2 enrichment

Prunella vulgaris was inoculated with different arbuscular mycorrhizal fungi (AMF) and grown at two concentrations of CO₂ (ambient, 350 μl l⁻¹, and elevated, 600 μl l⁻¹) to test whether a plants response to elevated CO₂ is dependent on the species of AMF colonizing the roots. Using compartments accessible only to AMF hyphae but not to roots, we also tested whether elevated CO₂ affects the growth of external AMF hyphae. Plant biomass was significantly greater at elevated than at ambient CO₂; the biomass of the root system, for example, increased by a factor of 2. The colonization of AMF inside the root remained constant, indicating that the total AMF inside the root system also increased by a factor of 2. The length of external AMF hyphae at elevated CO₂ was up to 5 times that at ambient CO₂, indicating that elevated CO₂ promoted allocation of AMF biomass to the external hyphae. The concentration and content of phosphorus in the stolons differed significantly between ambient and elevated CO₂ but this resulted in either an increase or a decrease, according to which AMF isolate occupied the roots. We hypothesized that an increase in external hyphal growth at elevated CO₂ would result in increased P acquistion by the plant. To test this we supplied phosphorus, in a compartment only accessible to AMF hyphae. Plants did not acquire more phosphorus at elevated CO₂ when phosphorus was added to this compartment. Large increases in AMF hyphal growth could, however, play a significant role in the movement of fixed carbon to the soil and increase soil aggregation. Received: 28 March 1998 / Accepted: 27 August 1998     

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

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

A meta-analysis of elevated CO2 effects on woody plant mass, form, and physiology

Quantitative integration of the literature on the effect of elevated CO₂ on woody plants is important to aid our understanding of forest health in coming decades and to better predict terrestrial feedbacks on the global carbon cycle. We used meta-analytic methods to summarize and interpret more than 500 reports of effects of elevated CO₂ on woody plant biomass accumulation and partitioning, gas exchange, and leaf nitrogen and starch content. The CO₂ effect size metric we used was the log-transformed ratio of elevated compared to ambient response means weighted by the inverse of the variance of the log ratio. Variation in effect size among studies was partitioned according to the presence of interacting stress factors, length of CO₂ exposure, functional group status, pot size, and type of CO₂ exposure facility. Both total biomass (W _T) and net CO₂ assimilation (A) increased significantly at about twice ambient CO₂, regardless of growth conditions. Low soil nutrient availability reduced the CO₂ stimulation of W _T by half, from +31% under optimal conditions to +16%, while low light increased the response to +52%. We found no significant shifts in biomass allocation under high CO₂. Interacting stress factors had no effect on the magnitude of responses of A to CO₂, although plants grown in growth chambers had significantly lower responses (+19%) than those grown in greenhouses or in open-top chambers (+54%). We found no consistent evidence for photosynthetic acclimation to CO₂ enrichment except in trees grown in pots <0.5 l (−36%) and no significant CO₂ effect on stomatal conductance. Both leaf dark respiration and leaf nitrogen were significantly reduced under elevated CO₂ (−18% and −16% respectively, data expressed on a leaf mass basis), while leaf starch content increased significantly except in low nutrient grown gymnosperms. Our results provide robust, statistically defensible estimates of elevated CO₂ effect sizes against which new results may be compared or for use in forest and climate model parameterization. Received: 16 May 1997 / Accepted: 9 September 1997     

Biomass allocation in old-field annual species grown in elevated CO2 environments: no evidence for optimal partitioning

Increased atmospheric carbon dioxide supply is predicted to alter plant growth and biomass allocation patterns. It is not clear whether changes in biomass allocation reflect optimal partitioning or whether they are a direct effect of increased growth rates. Plasticity in growth and biomass allocation patterns was investigated at two concentrations of CO₂ ([CO₂]) and at limiting and nonlimiting nutrient levels for four fast‐ growing old‐field annual species. Abutilon theophrasti, Amaranthus retroflexus, Chenopodium album, and Polygonum pensylvanicum were grown from seed in controlled growth chamber conditions at current (350 μmol mol^?1, ambient) and future‐ predicted (700 μmol mol^?1, elevated) CO₂ levels. Frequent harvests were used to determine growth and biomass allocation responses of these plants throughout vegetative development. Under nonlimiting nutrient conditions, whole plant growth was increased greatly under elevated [CO₂] for three C3 species and moderately increased for a C4 species (Amaranthus). No significant increases in whole plant growth were observed under limiting nutrient conditions. Plants grown in elevated [CO₂] had lower or unchanged root:shoot ratios, contrary to what would be expected by optimal partitioning theory. These differences disappeared when allometric plots of the same data were analysed, indicating that CO₂‐induced differences in root:shoot allocation were a consequence of accelerated growth and development rates. Allocation to leaf area was unaffected by atmospheric [CO₂] for these species. The general lack of biomass allocation responses to [CO₂] availability is in stark contrast with known responses of these species to light and nutrient gradients. We conclude that biomass allocation responses to elevated atmospheric [CO₂] are not consistent with optimal partitioning predictions.     

Towards an understanding of how phloem amino acid composition shapes elevated CO2‐induced changes in aphid population dynamics

1. The performance of foliage feeders tends to decrease under elevated CO₂, but the responses of phloem‐feeding insects have been much more equivocal. As phloem tissues are less accessible than whole‐plant tissues, much less is known about how phloem composition is altered under elevated CO₂ and the mechanisms driving changes in aphid performance. 2. In this study, the plant mechanisms underlying the performance of Rhopalosiphum padi aphids on Hordeum vulgare (barley) grown under ambient (390 ppm) and elevated (700 ppm) CO₂ were examined. We used aphid stylectomy to sample pure phloem from plants in CO₂‐controlled conditions and high‐performance liquid chromatography to analyse phloem samples for amino acid concentrations. 3. Aphid abundance significantly increased by 127% under elevated CO₂. Consequently, plant biomass decreased under elevated CO₂ in trials with herbivores present, possibly due to the increased herbivore load, but increased when aphids were absent. The intrinsic rate of population increase (r_m) was significantly higher under elevated CO₂; however, there were no statistically significant effects on aphid fecundity or development time. The concentration of individual amino acids tended to increase, although these increases were statistically significant in only a few cases. A principal components analysis revealed that the relative abundance (mol %) of those amino acids considered essential for aphids tended to increase under elevated CO₂. 4. These results indicate that CO₂ may affect nutrient translocation in plants in ways that are contrary to predictions about nitrogen metabolite responses to CO₂. Such plant biochemical responses may underlie observations of improved phloem feeder performance under elevated CO₂.     

Increased soil moisture content increases plant N uptake and the abundance of 15N in plant biomass

The natural abundance of ¹⁵N in plant biomass has been used to infer how N dynamics change with elevated atmospheric CO₂ and changing water availability. However, it remains unclear if atmospheric CO₂ effects on plant biomass ¹⁵N are driven by CO₂-induced changes in soil moisture. We tested whether ¹⁵N abundance (expressed as δ¹⁵N) in plant biomass would increase with increasing soil moisture content at two atmospheric CO₂ levels. In a greenhouse experiment we grew sunflower (Helianthus annuus) at ambient and elevated CO₂ (760 ppm) with three soil moisture levels maintained at 45, 65, and 85% of field capacity, thereby eliminating potential CO₂-induced soil moisture effects. The δ¹⁵N value of total plant biomass increased significantly with increased soil moisture content at both CO₂ levels, possibly due to increased uptake of ¹⁵N-rich organic N. Although not adequately replicated, plant biomass δ¹⁵N was lower under elevated than under ambient CO₂ after adjusting for plant N uptake effects. Thus, increases in soil moisture can increase plant biomass δ¹⁵N, while elevated CO₂ can decrease plant biomass δ¹⁵N other than by modifying soil moisture.     

Regenerating temperate forest mesocosms in elevated CO2: belowground growth and nitrogen cycling

The response of temperate forest ecosystems to elevated atmospheric CO₂ concentrations is important because these ecosystems represent a significant component of the global carbon cycle. Two important but not well understood processes which elevated CO₂ may substantially alter in these systems are regeneration and nitrogen cycling. If elevated CO₂ leads to changes in species composition in regenerating forest communities then the structure and function of these ecosystems may be affected. In most temperate forests, nitrogen appears to be a limiting nutrient. If elevated CO₂ leads to reductions in nitrogen cycling through increased sequestration of nitrogen in plant biomass or reductions in mineralization rates, long-term forest productivity may be constrained. To study these processes, we established mesocosms of regenerating forest communities in controlled environments maintained at either ambient (375 ppm) or elevated (700 ppm) CO₂ concentrations. Mesocosms were constructed from intact monoliths of organic forest soil. We maintained these mesocosms for 2 years without any external inputs of nitrogen and allowed the plants naturally present as seeds and rhizomes to regenerate. We used ¹⁵N pool dilution techniques to quantify nitrogen fluxes within the mesocosms at the end of the 2 years. Elevated atmospheric CO₂ concentration significantly affected a number of plant and soil processes in the experimental regenerating forest mesocosms. These changes included increases in total plant biomass production, plant C/N ratios, ectomycorrhizal colonization of tree fine roots, changes in tree fine root architecture, and decreases in plant NH₄ ⁺ uptake rates, gross NH₄ ⁺ mineralization rates, and gross NH₄ ⁺ consumption rates. In addition, there was a shift in the relative biomass contribution of the two dominant regenerating tree species; the proportion of total biomass contributed by white birch (Betula papyrifera) decreased and the proportion of total biomass contributed by yellow birch (B. alleghaniensis) increased. However, elevated CO₂ had no significant effect on the total amount of nitrogen in plant and soil microbial biomass. In this study we observed a suite of effects due to elevated CO₂, some of which could lead to increases in potential long term growth responses to elevated CO₂, other to decreases. The reduced plant NH₄ ⁺ uptake rates we observed are consistent with reduced NH₄ ⁺ availability due to reduced gross mineralization rates. Reduced NH₄ ⁺ mineralization rates are consistent with the increases in C/N ratios we observed for leaf and fine root material. Together, these data suggest the positive increases in plant root architectural parameters and mycorrhizal colonization may not be as important as the potential negative effects of reduced nitrogen availability through decreased decomposition rates in a future atmosphere with elevated CO₂. Received: 10 January 1997 / Accepted: 25 July 1997     

The fraction of expanding to expanded leaves determines the biomass response of Populus to elevated CO2

We examined whether the effects of elevated CO₂ on growth of 1-year old Populus deltoides saplings was a function of the assimilation responses of particular leaf developmental stages. Saplings were grown for 100 days at ambient (approximately 350 ppm) and elevated (ambient + 200 ppm) CO₂ in forced-air greenhouses. Biomass, biomass distribution, growth rates, and leaf initiation and expansion rates were unaffected by elevated CO₂. Leaf nitrogen (N), the leaf C:N ratio, and leaf lignin concentrations were also unaffected. Carbon gain was significantly greater in expanding leaves of saplings grown at elevated compared to ambient CO₂. The Rubisco content in expanding leaves was not affected by CO₂ concentration. Carbon gain and Rubisco content were significantly lower in fully expanded leaves of saplings grown at elevated compared to ambient CO₂, indicating CO₂-induced down-regulation in fully expanded leaves. Elevated CO₂ likely had no overall effect on biomass accumulation due to the more rapid decline in carbon gain as leaves matured in saplings grown at elevated compared to ambient CO₂. This decline in carbon gain has been documented in other species and shown to be related to a balance between sink/source balance and acclimation. Our data suggest that variation in growth responses to elevated CO₂ can result from differences in leaf assimilation responses in expanding versus expanded leaves as they develop under elevated CO₂. Received: 28 September 1998 / Accepted: 23 June 1999     

The effect of heat waves,elevated [CO2] and low soil water availability on northern red oak (Quercus rubra L.) seedlings

The frequency and intensity of heat waves are predicted to increase. This study investigates whether heat waves would have the same impact as a constant increase in temperature with the same heat sum, and whether there would be any interactive effects of elevated [CO₂] and soil moisture content. We grew Quercus rubra seedlings in treatment chambers maintained at either ambient or elevated [CO₂] (380 or 700 μmol CO₂ mol^?1) with temperature treatments of ambient, ambient +3 °C, moderate heat wave (+6 °C every other week) or severe heat wave (+12 °C every fourth week) temperatures. Averaged over a 4‐week period, and the entire growing season, the three elevated temperature treatments had the same average temperature and heat sum. Half the seedlings were watered to a soil water content near field capacity, half to about 50% of this value. Foliar gas exchange measurements were performed morning and afternoon (9:00 and 15:00 hours) before, during and after an applied heat wave in August 2010. Biomass accumulation was measured after five heat wave cycles. Under ambient [CO₂] and well‐watered conditions, biomass accumulation was highest in the +3 °C treatment, intermediate in the +6 °C heat wave and lowest in the +12 °C heat wave treatment. This response was mitigated by elevated [CO₂]. Low soil moisture significantly decreased net photosynthesis (A_net) and biomass in all [CO₂] and temperature treatments. The +12 °C heat wave reduced afternoon A_net by 23% in ambient [CO₂]. Although this reduction was relatively greater under elevated [CO₂], A_net values during this heat wave were still 34% higher than under ambient [CO₂]. We concluded that heat waves affected biomass growth differently than the same amount of heat applied uniformly over the growing season, and that the plant response to heat waves also depends on [CO₂] and soil moisture conditions.     

Elevated CO2 shifts the focus of tobacco plant defences from cucumber mosaic virus to the green peach aphid

Elevation in CO₂ concentration broadly impacts plant physiological characteristics, which influences herbivores and biotrophic pathogens, which in turn regulate the plant defensive response. In this study, responses of tobacco plants to stress in the form of the green peach aphid, Myzus persicae (Sulzer), or cucumber mosaic virus (CMV), or both aphid and CMV combined were investigated in open‐top chambers under ambient and elevated CO₂ concentrations. We measured aboveground biomass and foliar chlorophyll, nitrogen, non‐structural carbohydrates, soluble protein, total amino acid and nicotine content in tobacco plants and also measured aphid population dynamics, body weight, honeydew production and anti‐oxidative enzyme activities in individual aphids. Plants produced more secondary metabolites for defence in both CO₂ treatments when treated with aphid and CMV combined than with either alone. Aphid density significantly increased on CMV‐infected tobacco plants (relative to uninfected plants) under ambient CO₂ but not under elevated CO₂. This suggests that plant defences against virus and aphid would be more efficient under elevated CO₂. Plant defence appears to shift from plant virus to aphid under increasing CO₂ levels, which highlights the potential influences of multiple biotic stressors on plants under elevated CO₂.     

Elevated CO<Subscript>2</Subscript> Concentration and Drought Stress Exert Opposite Effects on Plant Biomass,Nitrogen, and Phosphorus Allocation in <Emphasis Type="Italic">Bothriochloa ischaemum</Emphasis>

Increased concentrations of atmospheric carbon dioxide (CO₂) and drought stress have greatly influenced plant growth, the status of nitrogen (N) and phosphorus (P), and N:P ratios. We identified the plant biomass, N and P distributional patterns, and N:P stoichiometry of a grass species on the Loess Plateau in China under elevated CO₂ concentration and drought stress conditions. Bothriochloa ischaemum, a C4 perennial herbaceous grass, was grown in pots at CO₂ concentrations of 400 (ambient) and 800 (elevated) μmol mol^?1 and at 60 ± 5 and 40 ± 5 % of field capacity. The elevated CO₂ concentration significantly increased plant total biomass, N concentration, N and P content, allocation of biomass to roots, and allocation of N to shoots, and increased the N:P ratios of whole plants and the shoots, especially under well-watered conditions. Drought stress significantly decreased plant biomass and plant N and P content, especially under elevated CO₂. Drought stress decreased the N:P ratios, but was only significant in the roots under ambient CO₂. Drought stress may attenuate the stimulation of plant growth and N and P acquisition by CO₂ enrichment, and projected elevated CO₂ concentrations may partially offset the negative effects of increased drought by increasing the assimilation of N and P.     

Near-term impacts of elevated CO2, nitrogen and fungal endophyte-infection on Lolium perenne L. growth, chemical composition and alkaloid production

Carbon dioxide has been rapidly accumulating in the atmosphere and is expected to continue to do so. This accumulation is presumed to have important direct effects on plant growth. The interacting affects of a small increase in CO₂ concentration (466 p.p.m., approximately 30% increase from current ambient conditions), nitrogen fertilization and fungal endophyte (Neotyphodium lolii) infection on the growth and chemical composition of perennial ryegrass (Lolium perenne) were investigated. It was found that dry mass production was approximately 50% greater under elevated CO₂ than under ambient CO₂, but only in conditions of high soil N. High molecular weight carbohydrates and total carbohydrates (LMW + HMW CHO) depended on an interaction between CO₂ and endophyte infection. Infected plants contained significantly more carbohydrate than endophyte-free plants, and the difference was greatest in ambient CO₂ conditions. Protein concentrations were also influenced by the interaction between CO₂ and endophyte-infection. Endophyte-free plants had 40% lower concentrations of soluble protein under elevated CO₂ than under ambient CO₂, but this CO₂ effect on soluble protein was largely absent in endophyte-infected plants. CO₂, endophyte-infection and nitrogen interacted to influence the total chlorophyll concentration of the grass such that chlorophyll concentration was always lower in elevated CO₂ but this decline was much greater in endophyte-free plants, particularly in conditions of high soil N. In the endophyte-infected plants, the concentrations of the pyrrolopyrazine alkaloid peramine depended on the interaction between CO₂ and N fertilization such that peramine concentrations declined with increasing N at ambient CO₂ but remained roughly constant across N levels at elevated CO₂. A similar pattern was seen for the ergot alkaloid ergovaline. The biochemical responses of perennial ryegrass to elevated CO₂ are clearly modified by the presence of endophytic fungi.