Acquisition and within-plant allocation of 13C and 15N in CO2-enriched Quercus robur plants

We assessed the effects of doubling atmospheric CO₂ concentration, [CO₂], on C and N allocation within pedunculate oak plants (Quercus robur L.) grown in containers under optimal water supply. A short-term dual ¹³CO₂ and ¹⁵NO₃^? labelling experiment was carried out when the plants had formed their third growing flush. The 22-week exposure to 700 μl l^?1 [CO₂] stimulated plant growth and biomass accumulation (+53% as compared with the 350 μl l^?1 [CO₂] treatment) but decreased the root/shoot biomass ratio (-23%) and specific leaf area (-18%). Moreover, there was an increase in net CO₂ assimilation rate (+37% on a leaf dry weight basis; +71% on a leaf area basis), and a decrease in both above- and below-ground CO₂ respiration rates (-32 and -26%, respectively, on a dry mass basis) under elevated [CO₂]. ¹³C acquisition, expressed on a plant mass basis or on a plant leaf area basis, was also markedly stimulated under elevated [CO₂] both after the 12-h ¹³CO₂ pulse phase and after the 60-h chase phase. Plant N content was increased under elevated CO₂ (+36%), but not enough to compensate for the increase in plant C content (+53%). Thus, the plant C/N ratio was increased (+13%) and plant N concentration was decreased (-11%). There was no effect of elevated [CO₂] on fine root-specific ¹⁵N uptake (amount of recently assimilated ¹⁵N per unit fine root dry mass), suggesting that modifications of plant N pools were merely linked to root size and not to root function. N concentration was decreased in the leaves of the first and second growing flushes and in the coarse roots, whereas it was unaffected by [CO₂] in the stem and in the actively growing organs (fine roots and leaves of the third growth flush). Furthermore, leaf N content per unit area was unaffected by [CO₂]. These results are consistent with the short-term optimization of N distribution within the plants with respect to growth and photosynthesis. Such an optimization might be achieved at the expense of the N pools in storage compartments (coarse roots, leaves of the first and second growth flushes). After the 60-h ¹³C chase phase, leaves of the first and second growth flushes were almost completely depleted in recent ¹³C under ambient [CO₂], whereas these leaves retained important amounts of recently assimilated ¹³C (carbohydrate reserves?) under elevated [CO₂].     

Effects of enhanced atmospheric CO2 and nutrient supply on the quality and subsequent decomposition of fine roots of Betula pendula Roth. and Picea sitchensis (Bong.) Carr.

Fine root litter derived from birch (Betula pendula Roth.) and Sitka spruce (Picea sitchensis (Bong.) Carr.) plants grown under two CO₂ atmospheric concentrations (350 ppm and 600 ppm) and two nutrient regimes was used for decomposition studies in laboratory microcosms. Although there were interactions between litter type, CO₂/fertiliser treatments and decomposition rates, in general, an increase in the C/N ratio of the root tissue was observed for roots of both species grown under elevated CO₂ in unfertilized soil. Both weight loss and respiration of decomposing birch roots were significantly reduced in materials derived from enriched CO₂, whilst the decomposition of spruce roots showed no such effect. A parallel experiment was performed using Betula pendula root litter grown under different N regimes, in order to test the relationship between C/N ratio of litter and root decomposition rate. A highly significant (p<0.001) negative correlation between C/N ratio and root litter respiration was found, with an r²=0.97. The results suggest that the increased C/N ratio of plant tissues induced by elevated CO₂ can result in a reduction of decomposition rate, with a resulting increase in forest soil C stores.     

Interactions between plant growth and soil nutrient cycling under elevated CO2: a meta-analysis

free air carbon dioxide enrichment (FACE) and open top chamber (OTC) studies are valuable tools for evaluating the impact of elevated atmospheric CO₂ on nutrient cycling in terrestrial ecosystems. Using meta‐analytic techniques, we summarized the results of 117 studies on plant biomass production, soil organic matter dynamics and biological N₂ fixation in FACE and OTC experiments. The objective of the analysis was to determine whether elevated CO₂ alters nutrient cycling between plants and soil and if so, what the implications are for soil carbon (C) sequestration. Elevated CO₂ stimulated gross N immobilization by 22%, whereas gross and net N mineralization rates remained unaffected. In addition, the soil C : N ratio and microbial N contents increased under elevated CO₂ by 3.8% and 5.8%, respectively. Microbial C contents and soil respiration increased by 7.1% and 17.7%, respectively. Despite the stimulation of microbial activity, soil C input still caused soil C contents to increase by 1.2% yr^?1. Namely, elevated CO₂ stimulated overall above‐ and belowground plant biomass by 21.5% and 28.3%, respectively, thereby outweighing the increase in CO₂ respiration. In addition, when comparing experiments under both low and high N availability, soil C contents (+2.2% yr^?1) and above‐ and belowground plant growth (+20.1% and+33.7%) only increased under elevated CO₂ in experiments receiving the high N treatments. Under low N availability, above‐ and belowground plant growth increased by only 8.8% and 14.6%, and soil C contents did not increase. Nitrogen fixation was stimulated by elevated CO₂ only when additional nutrients were supplied. These results suggest that the main driver of soil C sequestration is soil C input through plant growth, which is strongly controlled by nutrient availability. In unfertilized ecosystems, microbial N immobilization enhances acclimation of plant growth to elevated CO₂ in the long‐term. Therefore, increased soil C input and soil C sequestration under elevated CO₂ can only be sustained in the long‐term when additional nutrients are supplied.     

Combined effects of atmospheric CO<Subscript>2</Subscript> and N availability on the belowground carbon and nitrogen dynamics of aspen mesocosms

It is uncertain whether elevated atmospheric CO₂ will increase C storage in terrestrial ecosystems without concomitant increases in plant access to N. Elevated CO₂ may alter microbial activities that regulate soil N availability by changing the amount or composition of organic substrates produced by roots. Our objective was to determine the potential for elevated CO₂ to change N availability in an experimental plant-soil system by affecting the acquisition of root-derived C by soil microbes. We grew Populus tremuloides (trembling aspen) cuttings for 2 years under two levels of atmospheric CO₂ (36.7 and 71.5 Pa) and at two levels of soil N (210 and 970 μg N g^–1). Ambient and twice-ambient CO₂ concentrations were applied using open-top chambers, and soil N availability was manipulated by mixing soils differing in organic N content. From June to October of the second growing season, we measured midday rates of soil respiration. In August, we pulse-labeled plants with ¹⁴CO₂ and measured soil ¹⁴CO₂ respiration and the ¹⁴C contents of plants, soils, and microorganisms after a 6-day chase period. In conjunction with the August radio-labeling and again in October, we used ¹⁵N pool dilution techniques to measure in situ rates of gross N mineralization, N immobilization by microbes, and plant N uptake. At both levels of soil N availability, elevated CO₂ significantly increased whole-plant and root biomass, and marginally increased whole-plant N capital. Significant increases in soil respiration were closely linked to increases in root biomass under elevated CO₂. CO₂ enrichment had no significant effect on the allometric distribution of biomass or ¹⁴C among plant components, total ¹⁴C allocation belowground, or cumulative (6-day) ¹⁴CO₂ soil respiration. Elevated CO₂ significantly increased microbial ¹⁴C contents, indicating greater availability of microbial substrates derived from roots. The near doubling of microbial ¹⁴C contents at elevated CO₂ was a relatively small quantitative change in the belowground C cycle of our experimental system, but represents an ecologically significant effect on the dynamics of microbial growth. Rates of plant N uptake during both 6-day periods in August and October were significantly greater at elevated CO₂, and were closely related to fine-root biomass. Gross N mineralization was not affected by elevated CO₂. Despite significantly greater rates of N immobilization under elevated CO₂, standing pools of microbial N were not affected by elevated CO₂, suggesting that N was cycling through microbes more rapidly. Our results contained elements of both positive and negative feedback hypotheses, and may be most relevant to young, aggrading ecosystems, where soil resources are not yet fully exploited by plant roots. If the turnover of microbial N increases, higher rates of N immobilization may not decrease N availability to plants under elevated CO₂. Received: 12 February 1999 / Accepted: 2 March 2000     

The effects of elevated [CO2] on the C:N and C:P mass ratios of plant tissues

The influence of elevated CO₂ concentration ([CO₂]) during plant growth on the carbon:nutrient ratios of tissues depends in part on the time and space scales considered. Most evidence relates to individual plants examined over weeks to just a few years. The C:N ratio of live tissues is found to increase, decrease or remain the same under elevated [CO₂]. On average it increases by about 15% under a doubled [CO₂]. A testable hypothesis is proposed to explain why it increases in some situations and decreases in others. It includes the notion that only in the intermediate range of N-availability will C:N of live tissues increase under elevated [CO₂]. Five hypotheses to explain the mechanism of such increase in C:N are discussed; none of these options explains all the published results. Where elevated [CO₂] did increase the C:N of green leaves, that response was not necessarily expressed as a higher C:N of senesced leaves. An hypothesis is explored to explain the observed range in the degree of propogation of a CO₂ effect on live tissues through to the litter derived from them. Data on C:P ratios under elevated [CO₂] are sparse and also variable. They do not yet suggest a generalising-hypothesis of responses. Although, unlike for C:N, there is no theoretical expectation that C:P of plants would increase under elevated [CO₂], the average trend in the data is of such an increase. The processes determining the C:P response to elevated [CO₂] seem to be largely independent of those for C:N. Research to advance the topic should be structured to examine the components of the hypotheses to explain effects on C:N. This involves experiments in which plants are grown over the full range of N and of P availability from extreme limitation to beyond saturation. Measurements need to: distinguish structural from non-structural dry matter; organic from inorganic forms of the nutrient in the tissues; involve all parts of the plant to evaluate nutrient and C allocation changes with treatments; determine resorption factors during tissue senescence; and be made with cognisance of the temporal and spatial aspects of the phenomena involved. This revised version was published online in June 2006 with corrections to the Cover Date.     

Plant and microbial N acquisition under elevated atmospheric CO2 in two mesocosm experiments with annual grasses

The impact of elevated CO₂ on terrestrial ecosystem C balance, both in sign or magnitude, is not clear because the resulting alterations in C input, plant nutrient demand and water use efficiency often have contrasting impacts on microbial decomposition processes. One major source of uncertainty stems from the impact of elevated CO₂ on N availability to plants and microbes. We examined the effects of atmospheric CO₂ enrichment (ambient+370 μmol mol^?1) on plant and microbial N acquisition in two different mesocosm experiments, using model plant species of annual grasses of Avena barbata and A. fatua, respectively. The A. barbata experiment was conducted in a N‐poor sandy loam and the A. fatua experiment was on a N‐rich clayey loam. Plant–microbial N partitioning was examined through determining the distribution of a ¹⁵N tracer. In the A. barbata experiment, ¹⁵N tracer was introduced to a field labeling experiment in the previous year so that ¹⁵N predominantly existed in nonextractable soil pools. In the A. fatua experiment, ¹⁵N was introduced in a mineral solution [(¹⁵NH₄)₂SO₄ solution] during the growing season of A. fatua. Results of both N budget and ¹⁵N tracer analyses indicated that elevated CO₂ increased plant N acquisition from the soil. In the A. barbata experiment, elevated CO₂ increased plant biomass N by ca. 10% but there was no corresponding decrease in soil extractable N, suggesting that plants might have obtained N from the nonextractable organic N pool because of enhanced microbial activity. In the A. fatua experiment, however, the CO₂‐led increase in plant biomass N was statistically equal to the reduction in soil extractable N. Although atmospheric CO₂ enrichment enhanced microbial biomass C under A. barbata or microbial activity (respiration) under A. fatua, it had no significant effect on microbial biomass N in either experiment. Elevated CO₂ increased the colonization of A. fatua roots by arbuscular mycorrhizal fungi, which coincided with the enhancement of plant competitiveness for soluble soil N. Together, these results suggest that elevated CO₂ may tighten N cycling through facilitating plant N acquisition. However, it is unknown to what degree results from these short‐term microcosm experiments can be extrapolated to field conditions. Long‐term studies in less‐disturbed soils are needed to determine whether CO₂‐enhancement of plant N acquisition can significantly relieve N limitation over plant growth in an elevated CO₂ environment.     

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.     

Canopy N and P dynamics of a southeastern US pine forest under elevated CO2

Forest production is strongly nutrient limited throughout the southeastern US. If nutrient limitations constrain plant acquisition of essential resources under elevated CO₂, reductions in the mass or nutrient content of forest canopies could constrain C assimilation from the atmosphere. We tested this idea by quantifying canopy biomass, foliar concentrations of N and P, and the total quantity of N and P in a loblolly pine (Pinus taeda) canopy subject to 4 years of free-air CO₂ enrichment. We also used N:P ratios to detect N versus P limitation to primary production under elevated CO₂. Canopy biomass was significantly higher under elevated CO₂ during the first 4 years of this experiment. Elevated CO₂ significantly reduced the concentration of N in loblolly pine foliage (5% relative to ambient CO₂) but not P. Despite the slight reduction foliage N concentrations, there were significant increases in canopy N and P contents under elevated CO₂. Foliar N:P ratios were not altered by elevated CO₂ and were within a range suggesting forest production is N limited not P limited. Despite the clear limitation of NPP by N under ambient and elevated CO₂ at this site, there is no evidence that the mass of N or P in the canopy is declining through the first 4 years of CO₂ fumigation. As a consequence, whole-canopy C assimilation is strongly stimulated by elevated CO₂ making this forest a larger net C sink under elevated CO₂ than under ambient CO₂. We discuss the potential for future decreases in canopy nutrient content as a result of limited changes in the size of the plant-available pools of N under elevated CO₂.     

Elevated CO2 and plant nitrogen-use: is reduced tissue nitrogen concentration size-dependent?

Plants often respond to elevated atmospheric CO₂ levels with reduced tissue nitrogen concentrations relative to ambient CO₂-grown plants when comparisons are made at a common time. Another common response to enriched CO₂ atmospheres is an acceleration in plant growth rates. Because plant nitrogen concentrations are often highest in seedlings and subsequently decrease during growth, comparisons between ambient and elevated CO₂-grown plants made at a common time may not demonstrate CO₂-induced reductions in plant nitrogen concentration per se. Rather, this comparison may be highlighting differences in nitrogen concentration between bigger, more developed plants and smaller, less developed plants. In this study, we directly examined whether elevated CO₂ environments reduce plant nitrogen concentrations independent of changes in plant growth rates. We grew two annual plant species. Abutilon theophrasti (C₃ photosynthetic pathway) and Amaranthus retroflexus (C₄ photosynthetic pathway), from seed in glass-sided growth chambers with atmospheric CO₂ levels of 350 mol·mol^–1 or 700 mol·mol^–1 and with high or low fertilizer applications. Individual plants were harvested every 2 days starting 3 days after germination to determine plant biomass and nitrogen concentration. We found: 1. High CO₂-grown plants had reduced nitrogen concentrations and increased biomass relative to ambient CO₂-grown plants when compared at a common time; 2. Tissue nitrogen concentrations did not vary as a function of CO₂ level when plants were compared at a common size; and 3. The rate of biomass accumulation per rate of increase in plant nitrogen was unaffected by CO₂ availability, but was altered by nutrient availability. These results indicate that a CO₂-induced reduction in plant nitrogen concentration may not be due to physiological changes in plant nitrogen use efficiency, but is probably a size-dependent phenomenon resulting from accelerated plant growth.     

Leaf litter production and decomposition in a poplar short-rotation coppice exposed to free air CO2 enrichment (POPFACE)

The capacity of forest ecosystems to sequester C in the soil relies on the net balance between litter production above, as well as, below ground, and decomposition processes. Nitrogen mineralization and its availability for plant growth and microbial activity often control the speed of both processes. Litter production, decomposition and N mineralization are strongly interdependent. Thus, their responses to global environmental changes (i.e. elevated CO₂, climate, N deposition, etc.) cannot be fully understood if they are studied in isolation. In the present experiment, we investigated litter fall, litter decomposition and N dynamics in decomposing litter of three Populus spp., in the second and third growing season of a short rotation coppice under FACE. Elevated CO₂ did not affect annual litter production but slightly retarded litter fall in the third growing season. In all species, elevated CO₂ lowered N concentration, resulting in a reduction of N input to the soil via litter fall, but did not affect lignin concentrations. Litter decomposition was studied in bags incubated in situ both in control and FACE plots. Litter lost between 15% and 18% of the original mass during the eight months of field incubation. On average, litter produced under elevated CO₂ attained higher residual mass than control litter. On the other end, when litter was incubated in FACE plots it exhibited higher decay rates. These responses were strongly species‐specific. All litter increased their N content during decomposition, indicating immobilization of N from external sources. Independent of the initial quality, litter incubated on FACE soils immobilized less N, possibly as a result of lower N availability in the soil. Indeed, our results refer to a short‐term decomposition experiment. However, according to a longer‐term model extrapolation of our results, we anticipate that in Mediterranean climate, under elevated atmospheric CO₂, soil organic C pool of forest ecosystems may initially display faster turnover, but soil N availability will eventually limit the process.     

Elevated CO2 and temperature alter nitrogen allocation in Douglas-fir

The effects of elevated CO₂ and temperature on principal carbon constituents (PCC) and C and N allocation between needle, woody (stem and branches) and root tissue of Pseudotsuga menziesii Mirb. Franco seedlings were determined. The seedlings were grown in sun‐lit controlled‐environment chambers that contained a native soil. Chambers were controlled to reproduce ambient or ambient +180 ppm CO₂ and either ambient temperature or ambient +3.5 °C for 4 years. There were no significant CO₂ × temperature interactions; consequently the data are presented for the CO₂ and temperature effects. At the final harvest, elevated CO₂ decreased the nonpolar fraction of the PCC and increased the polar fraction and amount of sugars in the needles. In contrast, elevated temperature increased the nonpolar fraction of the PCC and decreased sugars in needles. There were no CO₂ or temperature effects on the PCC fractions in the woody tissue or root tissue. Elevated CO₂ and temperature had no significant effects on the C content of any of the plant tissues or fractions. In contrast, the foliar N content declined under elevated CO₂ and increased under elevated temperature; there were no significant effects in other tissues. The changes in the foliar N concentrations were in the cellulose and lignin fractions, the fractions, which contain protein, and are the consequences of changes in N allocation under the treatments. These results indicate reallocation of N among plant organs to optimize C assimilation, which is mediated via changes in the selectivity of Rubisco and carbohydrate modulation of gene expression.     

Elevated CO2 stimulates associative N2 fixation in a C3 plant of the Chesapeake Bay wetland

In this study, the response of N₂ fixation to elevated CO₂ was measured in Scirpus olneyi, a C₃ sedge, and Spartina patens, a C₄ grass, using acetylene reduction assay and ¹⁵N₂ gas feeding. Field plants grown in PVC tubes (25 cm long, 10 cm internal diameter) were used. Exposure to elevated CO₂ significantly (P < 0·05) caused a 35% increase in nitrogenase activity and 73% increase in ¹⁵N incorporated by Scirpus olneyi. In Spartina patens, elevated CO₂ (660 ± 1 μ mol mol ^{− 1}) increased nitrogenase activity and ¹⁵N incorporation by 13 and 23%, respectively. Estimates showed that the rate of N₂ fixation in Scirpus olneyi under elevated CO₂ was 611 ± 75 ng ¹⁵N fixed plant ^{− 1} h ^{− 1} compared with 367 ± 46 ng ¹⁵N fixed plant ^{− 1} h ^{− 1} in ambient CO₂ plants. In Spartina patens, however, the rate of N₂ fixation was 12·5 ± 1·1 versus 9·8 ± 1·3 ng ¹⁵N fixed plant ^{− 1} h ^{− 1} for elevated and ambient CO₂, respectively. Heterotrophic non-symbiotic N₂ fixation in plant-free marsh sediment also increased significantly (P < 0·05) with elevated CO₂. The proportional increase in ¹⁵N₂ fixation correlated with the relative stimulation of photosynthesis, in that N₂ fixation was high in the C₃ plant in which photosynthesis was also high, and lower in the C₄ plant in which photosynthesis was relatively less stimulated by growth in elevated CO₂. These results are consistent with the hypothesis that carbon fixation in C₃ species, stimulated by rising CO₂, is likely to provide additional carbon to endophytic and below-ground microbial processes.     

Soil [N] modulates soil C cycling in CO2‐fumigated tree stands: a meta‐analysis

Under elevated atmospheric CO₂ concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO₂ effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta‐analysis to test the hypotheses that: (1) elevated atmospheric CO₂ stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO₂ induces a C allocation shift towards below‐ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO₂. Soil N concentration strongly interacted with CO₂ fumigation: the effect of elevated CO₂ on fine root biomass and –production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO₂ are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.     

Soil 13C–15N dynamics in an N2-fixing clover system under long-term exposure to elevated atmospheric CO2

Reduced soil N availability under elevated CO₂ may limit the plant's capacity to increase photosynthesis and thus the potential for increased soil C input. Plant productivity and soil C input should be less constrained by available soil N in an N₂‐fixing system. We studied the effects of Trifolium repens (an N₂‐fixing legume) and Lolium perenne on soil N and C sequestration in response to 9 years of elevated CO₂ under FACE conditions. ¹⁵N‐labeled fertilizer was applied at a rate of 140 and 560 kg N ha^?1 yr^?1 and the CO₂ concentration was increased to 60 Pa pCO₂ using ¹³C‐depleted CO₂. The total soil C content was unaffected by elevated CO₂, species and rate of ¹⁵N fertilization. However, under elevated CO₂, the total amount of newly sequestered soil C was significantly higher under T. repens than under L. perenne. The fraction of fertilizer‐N (f_N) of the total soil N pool was significantly lower under T. repens than under L. perenne. The rate of N fertilization, but not elevated CO₂, had a significant effect on f_N values of the total soil N pool. The fractions of newly sequestered C (f_C) differed strongly among intra‐aggregate soil organic matter fractions, but were unaffected by plant species and the rate of N fertilization. Under elevated CO₂, the ratio of fertilizer‐N per unit of new C decreased under T. repens compared with L. perenne. The L. perenne system sequestered more ¹⁵N fertilizer than T. repens: 179 vs. 101 kg N ha^?1 for the low rate of N fertilization and 393 vs. 319 kg N ha^?1 for the high N‐fertilization rate. As the loss of fertilizer‐¹⁵N contributed to the ¹⁵N‐isotope dilution under T. repens, the input of fixed N into the soil could not be estimated. Although N₂ fixation was an important source of N in the T. repens system, there was no significant increase in total soil C compared with a non‐N₂‐fixing L. perenne system. This suggests that N₂ fixation and the availability of N are not the main factors controlling soil C sequestration in a T. repens system.     

Interacting effects of elevated CO2, nutrient availability and plant species on a generalist invertebrate herbivore

By affecting plant growth and phytochemistry elevated CO₂ may have indirect effects on the performance of herbivores. These effects show considerable variability across studies and may depend on nutrient availability, the carbon/nutrient‐balance in plant tissues and the secondary metabolism of plants. We studied the responses to elevated CO₂ and different nutrient availability of 12 herbaceous plant species differing in their investment into secondary compounds. Caterpillars of the generalist herbivore Spodoptera littoralis were reared on the leaves produced and their consumption and growth rates analysed. Elevated CO₂ resulted in a similar increase of biomass in all plant species, whereas the positive effect of fertilization varied among plant species. Specific leaf weight was influenced by elevated CO₂, but the effect depended on nutrient level and identity of plant species. Elevated CO₂ increased the C/N ratio of the leaves of most species. Caterpillars consumed more leaf material when plants were grown under elevated CO₂ and low nutrients. This indicates compensatory feeding due to lower tissue quality. However, the effects of elevated CO₂, nutrient availability and plant species identity on leaf consumption interacted. Both the effects of CO₂ and nutrient availability on the relative growth rate of the herbivore depended on the plant species. The feeding rate of S. littoralis on plant species that do not produce nitrogen‐containing secondary compounds (NCSC) was higher under low nutrient availability. In contrast, in plants producing NCSC nutrient availability had no effect on the feeding rate. This suggests that compensatory feeding in response to low nutrient contents may not be possible if plants produce NCSC. We conclude that elevated CO₂ causes species‐specific changes in the quality of plant tissues and consequently in changes in the preferences of herbivores for plant species. This could result in changes in plant community composition.     

Root production and tissue quality in a shortgrass steppe exposed to elevated CO<Subscript>2</Subscript>: Using a new ingrowth method

A modified root ingrowth method was developed to minimize destructive sampling in experiments with limited space, and used to estimate belowground net primary production and root tissue quality in a native semiarid grassland exposed to elevated CO₂ for five years. Increases in root production of over 60% were observed with elevated CO₂ during years of intermediate levels of precipitation, with smaller effects in a very wet year and no effects in a very dry year. Aboveground to belowground production ratios, and the depth distribution of root production, did not differ between ambient and elevated CO₂ treatments. Root soluble concentrations increased an average of 11% and lignin concentrations decreased an average of 6% with elevated CO₂, while nitrogen concentrations decreased an average of 21%. However, most tissue quality responses to CO₂ varied greatly among years, and C:N ratios were higher in only one year (22 ambient vs. 33 elevated). Among years, root nitrogen concentrations declined with increasing aboveground plant nitrogen yield, and increased over the study period. Estimates of root production by the ingrowth donut method were much lower than previous estimates in the shortgrass steppe based on ¹⁴C decay. We discuss reasons why all ingrowth methods will always result in relative rather than absolute estimates of root production.     

Carbon and nitrogen pools and mineralization in a grassland gley soil under elevated carbon dioxide at a natural CO2 spring

The growth and chemical composition of most plants are influenced by elevated CO₂, but accompanying effects on soil organic matter pools and mineralization are less clearly defined, partly because of the short‐term nature of most studies. Herein we describe soil properties from a naturally occurring cold CO₂ spring (Hakanoa) in Northland, New Zealand, at which the surrounding vegetation has been exposed to elevated CO₂ for at least several decades. The mean annual temperature at this site is ≈ 15.5 °C and rainfall ≈ 1550 mm. The site was unfertilized and ungrazed, with a vegetation of mainly C3 and C4 grasses, and had moderate levels of ‘available’ P. Two soils were present ? a gley soil and an organic soil – but only the gley soil is examined here. Average atmospheric CO₂ concentrations at 17 sampling locations in the gley soil area ranged from 372 to 670 ppmv. In samples at 0–5 cm depth, pH averaged 5.4; average values for organic C were 150 g, total N 11 g, microbial C 3.50 g, and microbial N 0.65 g kg^?1, respectively. Under standardized moisture conditions at 25 °C, average rates of CO₂‐C production (7–14 days) were 5.4 mg kg^?1 h^?1 and of net mineral‐N production (14 ?42 days) 0.40 mg kg^?1 h^?1. These properties were all correlated positively and significantly (P < 0.10) with atmospheric CO₂ concentrations, but not with soil moisture (except for CO₂‐C production) or with clay content; they were, however, correlated negatively and mainly significantly with soil pH. In spite of uncertainties associated with the uncontrolled environment of naturally occurring springs, we conclude that storage of C and N can increase under prolonged exposure to elevated CO₂, and may include an appreciable labile fraction in mineral soil with an adequate nutrient supply.     

Effects of elevated CO2 on the protein concentration of food crops: a meta-analysis

Meta‐analysis techniques were used to examine the effect of elevated atmospheric carbon dioxide [CO₂] on the protein concentrations of major food crops, incorporating 228 experimental observations on barley, rice, wheat, soybean and potato. Each crop had lower protein concentrations when grown at elevated (540–958 μmol mol^?1) compared with ambient (315–400 μmol mol^?1) CO₂. For wheat, barley and rice, the reduction in grain protein concentration was ～10–15% of the value at ambient CO₂. For potato, the reduction in tuber protein concentration was 14%. For soybean, there was a much smaller, although statistically significant reduction of protein concentration of 1.4%. The magnitude of the CO₂ effect on wheat grains was smaller under high soil N conditions than under low soil N. Protein concentrations in potato tubers were reduced more for plants grown at high than at low concentrations of ozone. For soybean, the ozone effect was the reverse, as elevated CO₂ increased the protein concentration of soybean grown at high ozone concentrations. The magnitude of the CO₂ effect also varied depending on experimental methodology. For both wheat and soybean, studies performed in open‐top chambers produced a larger CO₂ effect than those performed using other types of experimental facilities. There was also indication of a possible pot artifact as, for both wheat and soybean, studies performed in open‐top chambers showed a significantly greater CO₂ effect when plants were rooted in pots rather than in the ground. Studies on wheat also showed a greater CO₂ effect when protein concentration was measured in whole grains rather than flour. While the magnitude of the effect of elevated CO₂ varied depending on the experimental procedures, a reduction in protein concentration was consistently found for most crops. These findings suggest that the increasing CO₂ concentrations of the 21st century are likely to decrease the protein concentration of many human plant foods.     

Effects of elevated atmospheric carbon dioxide on amino acid and NH4+-N cycling in a temperate pine ecosystem

Rising atmospheric carbon dioxide (CO₂) is expected to increase forest productivity, resulting in greater carbon (C) storage in forest ecosystems. Because elevated atmospheric CO₂ does not increase nitrogen (N) use efficiency in many forest tree species, additional N inputs will be required to sustain increased net primary productivity (NPP) under elevated atmospheric CO₂. We investigated the importance of free amino acids (AAs) as a source for forest N uptake at the Duke Forest Free Air CO₂ Enrichment (FACE) site, comparing its importance with that of better‐studied inorganic N sources. Potential proteolytic enzyme activity was monitored seasonally, and individual AA concentrations were measured in organic horizon extracts. Potential free AA production in soils ranged from 190 to 690 nmol N g⁻¹ h⁻¹ and was greater than potential rates of soil NH₄⁺ production. Because of this high potential rate of organic N production, we determined (1) whether intact AA uptake occurs by Pinus taeda L., the dominant tree species at the FACE site, (2) if the rate of cycling of AAs is comparable with that of ammonium (NH₄⁺), and (3) if atmospheric CO₂ concentration alters the aforementioned N cycling processes. A field experiment using universally labeled ammonium (¹⁵NH₄⁺) and alanine (¹³C₃H₇¹⁵NO₂) demonstrated that ¹⁵N is more readily taken up by plants and heterotrophic microorganisms as NH₄⁺. Pine roots and microbes take up on average 2.4 and two times as much NH₄^{+ 15}N compared with alanine ¹⁵N 1 week after tracer application. N cycling through soil pools was similar for alanine and NH₄⁺, with the greatest ¹⁵N tracer recovery in soil organic matter, followed by microbial biomass, dissolved organic N, extractable NH₄⁺, and fine roots. Stoichiometric analyses of ¹³C and ¹⁵N uptake demonstrated that both plants and soil microorganisms take up alanine directly, with a ¹³C : ¹⁵N ratio of 3.3 : 1 in fine roots and 1.5 : 1 in microbial biomass. Our results suggest that intact AA (alanine) uptake contributes substantially to plant N uptake in loblolly pine forests. However, we found no evidence supporting increased recovery of free AAs in fine roots under elevated CO₂, suggesting plants will need to acquire additional N via other mechanisms, such as increased root exploration or increased N use efficiency.     

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.