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.     

RETRACTED ARTICLE: CO<Subscript>2</Subscript> effects on plant nutrient concentration depend on plant functional group and available nitrogen: a meta-analysis

Elevated CO₂ is expected to lower plant nutrient concentrations via carbohydrate dilution and increased nutrient use efficiency. Elevated CO₂ consistently lowers plant foliar nitrogen, but there is no consensus on CO₂ effects across the range of plant nutrients. We used meta-analysis to quantify elevated CO₂ effects on leaf, stem, root, and seed concentrations of B, Ca, Cu, Fe, K, Mg, Mn, P, S, and Zn among four plant functional groups and two levels of N fertilization. CO₂ effects on plant nutrient concentration depended on the nutrient, plant group, tissue, and N status. CO₂ reduced B, Cu, Fe, and Mg, but increased Mn concentration in the leaves of N₂ fixers. Elevated CO₂ increased Cu, Fe, and Zn, but lowered Mn concentration in grass leaves. Tree leaf responses were strongly related to N status: CO₂ significantly decreased Cu, Fe, Mg, and S at high N, but only Fe at low N. Elevated CO₂ decreased Mg and Zn in crop leaves grown with high N, and Mn at low N. Nutrient concentrations in crop roots were not affected by CO₂ enrichment, but CO₂ decreased Ca, K, Mg and P in tree roots. Crop seeds had lower S under elevated CO₂. We also tested the validity of a “dilution model.” CO₂ reduced the concentration of plant nutrients 6.6% across nutrients and plant groups, but the reduction is less than expected (18.4%) from carbohydrate accumulation alone. We found that elevated CO₂ impacts plant nutrient status differently among the nutrient elements, plant functional groups, and among plant tissues. Our synthesis suggests that differences between plant groups and plant organs, N status, and differences in nutrient chemistry in soils preclude a universal hypothesis strictly related to carbohydrate dilution regarding plant nutrient response to elevated CO₂.     

Microbial Community Responses to Atmospheric Carbon Dioxide Enrichment in a Warm-Temperate Forest

Forest productivity depends on nutrient supply, and sustained increases in forest productivity under elevated carbon dioxide (CO₂) may ultimately depend on the response of microbial communities to changes in the quantity and chemistry of plant-derived substrates, We investigated microbial responses to elevated CO₂ in a warm-temperate forest under free-air CO₂ enrichment for 5 years (1997–2001). The experiment was conducted on three 30 m diameter plots under ambient CO₂ and three plots under elevated CO₂ (200 ppm above ambient). To understand how microbial processes changed under elevated CO₂, we assayed the activity of nine extracellular enzymes responsible for the decomposition of labile and recalcitrant carbon (C) substrates and the release of nitrogen (N) and phosphorus (P) from soil organic matter. Enzyme activities were measured three times per year in a surface organic horizon and in the top 15 cm of mineral soil. Initially, we found significant increases in the decomposition of labile C substrates in the mineral soil horizon under elevated CO₂; this overall pattern was present but much weaker in the O horizon. Beginning in the 4th year of this study, enzyme activities in the O horizon declined under elevated CO₂, whereas they continued to be stimulated in the mineral soil horizon. By year 5, the degradation of recalcitrant C substrates in mineral soils was significantly higher under elevated CO₂. Although there was little direct effect of elevated CO₂ on the activity of N- and P-releasing enzymes, the activity of nutrient-releasing enzymes relative to those responsible for C metabolism suggest that nutrient limitation is increasingly regulating microbial activity in the O horizon. Our results show that the metabolism of microbial communities is significantly altered by the response of primary producers to elevated CO₂. We hypothesize that ecosystem responses to elevated CO₂ are shifting from primary production to decomposition as a result of increasing nutrient limitation.     

Influence of vesicular arbuscular mycorrhizae and leaf age on net gas exchange of citrus leaves

The purpose of this study was to test the hypothesis that vesicular arbuscular mycorrhizal (VAM) fungi affect net assimilation of CO₂ (A) of different-aged citrus leaves independent of mineral nutrition effects of mycorrhizae. Citrus aurantium L., sour orange plants were grown for 6 months in a sandy soil low in phosphorus that was either infested with the VAM fungus, Glomus intraradices Schenck & Smith, or fertilized with additional phosphorus and left nonmycorrhizal (NM). Net CO₂ assimilation, stomatal conductance, water use efficiency, and mineral nutrient status for expanding, recently expanded, and mature leaves were evaluated as well as plant size and relative growth rate of leaves. Nutrient status and net gas exchange varied with leaf age. G. intraradices-inoculated plants had well-established colonization (79% of root length) and were comparable in relative growth rate and size at final harvest with NM plants. Leaf mineral concentrations were generally the same for VAM and NM plants except for nitrogen. Although leaf nitrogen was apparently sufficient for high rates of A, VAM plants did have higher nitrogen concentrations than NM at the time of gas exchange measurements. G. intraradices had no effect on A, stomatal conductance, or water use efficiency, irrespective of leaf age. These results show that well-established VAM colonization does not affect net gas exchange of citrus plants that are comparable in size, growth rate, and nutritional status with NM plants.     

Phosphorus supply and arbuscular mycorrhizas increase growth and net gas exchange responses of two Citrus spp. grown at elevated [CO2]

We hypothesized that greater photosynthate supply at elevated [CO₂] could compensate for increased below-ground C demands of arbuscular mycorrhizas. Therefore, we investigated plant growth, mineral nutrition, starch, and net gas exchange responses of two Citrus spp. to phosphorus (P) nutrition and mycorrhizas at elevated atmospheric [CO₂]. Half of the seedlings of sour orange (C. aurantium L.) and ‘Ridge Pineapple’ sweet orange (C. sinensis L. Osbeck) were inoculated with the arbuscular mycorrhizal (AM) fungus, Glomus intraradices Schenck and Smith and half were non-mycorrhizal (NM). Plants were grown at ambient or 2X ambient [CO₂] in unshaded greenhouses for 11 weeks and fertilized daily with nutrient solution either without added P or with 2 mM P in a low-P soil. High P supply reduced AM colonization whereas elevated [CO₂] counteracted the depressive effect of P on intraradical colonization and vesicle development. Seedlings grown at either elevated [CO₂], high P or with G. intraradices had greater growth, net assimilation of CO₂ (A _CO2) in leaves, leaf water-use efficiency, leaf dry wt/area, leaf starch and carbon/nitrogen (C/N) ratio. Root/whole plant dry wt ratio was decreased by elevated [CO₂], P, and AM colonization. Mycorrhizal seedlings had higher leaf-P status but lower leaf N and K concentrations than nonmycorrhizal seedlings which was due to growth dilution effects. Starch in fibrous roots was increased by elevated [CO₂] but reduced by G. intraradices, especially at low-P supply. In fibrous roots, elevated [CO₂] had no effect on C/N, but AM colonization decreased C/N in both Citrus spp. grown at low-P supply. Overall, there were no species differences in growth or A _CO2. Mycorrhizas did not increase plant growth at ambient [CO₂]. At elevated [CO₂], however, mycorrhizas stimulated growth at both P levels in sour orange, the more mycorrhiza-dependent species, but only at low-P in sweet orange, the less dependent species. At low-P and elevated [CO₂], colonization by the AM fungus increased A _CO2 in both species but more so in sour orange than in sweet orange. Leaf P and root N concentrations were increased more and root starch level was decreased less by AM in sour orange than in sweet orange. Thus, the additional [CO₂] availability to mycorrhizal plants increased CO₂ assimilation, growth and nutrient uptake over that of NM plants especially in sour orange under P limitation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of CO2 and nutrient enrichment on tissue quality of two California annuals

The effects of CO₂ enrichment and soil nutrient status on tissue quality were investigated and related to the potential effect on growth and decomposition. Two California annuals, Avena fatua and Plantago erecta, were grown at ambient and ambient plus 35 Pa atmospheric CO₂ in nutrient unamended and amended serpentine soil. Elevated CO₂ led to significantly increased Avena shoot nitrogen concentrations in the nutrient amended treatment. It also led to decreased lignin concentrations in Avena roots in both nutrient treatments, and in Plantago shoots and roots with nutrient addition. Concentrations of total nonstructural carbohydrate (TNC) and carbon did not change with elevated CO₂ in either species. As a consequence of increased biomass accumulation, increased CO₂ led to larger total pools of TNC, lignin, total carbon, and total nitrogen in Avena with nutrient additions. Doubling CO₂ had no significant effect on Plantago. Given the limited changes in the compounds related to decomposibility and plant growth, effects of increased atmospheric CO₂ mediated through tissue composition on Avena and Plantago are likely to be minor and depend on site fertility. This study suggests that other factors such as litter moisture, whether or not litter is on the ground, and biomass allocation among roots and shoots, are likely to be more important in this California grassland ecosystem. CO₂ could influence those directly as well as indirectly.     

Ectomycorrhizal fungi associated with Pinus sylvestris seedlings respond differently to increased carbon and nitrogen availability: implications for ecosystem responses to global change

The ectomycorrhizal (ECM) symbiosis can cause both positive and negative feedback with trees under elevated CO₂. Positive feedback arises if the additional carbon (C) increases both nutrient uptake by the fungus and nutrient transfer to the plant, whereas negative feedback results from increased nutrient uptake and immobilization by the fungus and reduced transfer to the plant. Because species of ECM fungi differ in their C and nitrogen (N) demand, understanding fungal species‐specific responses to variation in C and N supply is essential to predict impacts of global change. We investigated fungal species‐specific responses of ECM Scots pine (Pinus sylvestris) seedlings under ambient and elevated CO₂ (350 or 700 μL L⁻¹ CO₂) and under low and high mineral N availability. Each seedling was associated with one of the following ECM species: Hebeloma cylindrosporum, Laccaria bicolor and Suillus bovinus. The experiment lasted 103 days. During the final 27 days, seedlings were labeled with ¹⁴CO₂ and ¹⁵N. Most plant and fungal parameters were significantly affected by fungal species, CO₂ level and N supply. Interactions between fungal species and CO₂ were also regularly significant. At low N availability, elevated CO₂ had the smallest impact on the photosynthetic performance of seedlings inoculated with H. cylindrosporum and the largest impact on seedlings with S. bovinus. At ambient CO₂, increasing N supply had the smallest impact on seedlings inoculated with S. bovinus and the largest on seedlings inoculated with H. cylindrosporum. At low N availability, extraradical hyphal length increased after doubling CO₂ level, but this was significant only for L. bicolor. At ambient CO₂, increasing N levels reduced hyphal length for both H. cylindrosporum and S. bovinus, but not for L. bicolor. We discuss the potential interplay of two major elements of global change, elevated CO₂ and increased N availability, and their effects on plant growth. We conclude that increased N supply potentially relieves mycorrhiza‐induced progressive N limitation under elevated CO₂.     

Atmospheric CO2, plant nitrogen status and the susceptibility of plants to an acute increase in temperature

Abstract. Elevated levels of CO₂ in the atmosphere are expected to affect plant performance and may alter global temperature patterns. Changes in mean air temperatures that might be induced by rising levels of CO₂ and other greenhouse gases could also be accompanied by increased variability in daily temperatures such that acute increases in air temperature may be more likely than at present. Consequently, we investigated whether plants grown in a CO₂ enriched atmosphere would be differently affected by a heat shock than plants grown at ambient CO₂ levels. Plants of a C₃ annual (Abutilon theophrasti), a C₃ annual crop (Sinapis alba) and a C₄ annual (Amaranthus retroflexus) were grown from seed in growth chambers under either 400 or 700cm³ m^?3 CO₂, and were fertilized with either a high or low nutrient regime. Young seedlings of S. alba, as well as plants of all species in either the vegetative or reproductive phase of growth were exposed to a 4-h heat shock in which the temperature was raised an additional 14–23°C (depending on plant age). Total biomass and reproductive biomass were examined to determine the effect of CO₂, nutrient and heat shock treatments on plant performance. Heat shock, CO₂, and nutrient treatments, all had some significant effects on plant performance, but plants from both CO₂ treatments responded similarly to heat shocks. We also found, as expected, that plants grown under high CO₂ had dramatically decreased tissue N concentrations relative to plants grown under ambient conditions. We predicted that high-CO₂-grown plants would be more susceptible to a heat shock than ambient-CO₂-grown plants, because the reduced N concentrations of high-CO₂ grown plants could result in the reduced synthesis of heat shock proteins and reduced thermotolerance. Although we did not examine heat shock proteins, our results showed little relationship between plant nitrogen status and the ability of a plant to tolerate an acute increase in temperature.     

Spatial heterogeneity in soil nutrient supply modulates nutrient and biomass responses to multiple global change drivers in model grassland communities

Changes in the atmospheric concentration of carbon dioxide ([CO₂]), nutrient availability and biotic diversity are three major drivers of the ongoing global change impacting terrestrial ecosystems worldwide. While it is well established that soil nutrient heterogeneity exerts a strong influence on the development of plant individuals and communities, it is virtually unknown how nutrient heterogeneity and global change drivers interact to affect plant performance and ecosystem functioning. We conducted a microcosm experiment to evaluate the effect of simultaneous changes in [CO₂], nutrient heterogeneity (NH), nutrient availability (NA) and species evenness on the biomass and nutrient uptake patterns of assemblages formed by Lolium perenne, Plantago lanceolata and Holcus lanatus. When the nutrients were heterogeneously supplied, assemblages exhibited precise root foraging patterns, and had higher above‐ and belowground biomass (average increases of 32% and 29% for above‐ and belowground biomass, respectively). Nutrient heterogeneity also modulated the effects of NA on biomass production, complementarity in nitrogen uptake and below: aboveground ratio, as well as those of [CO₂] on the nutrient use efficiency at the assemblage level. Our results show that nutrient heterogeneity has the potential to influence the response of plant assemblages to simultaneous changes in [CO₂], nutrient availability and biotic diversity, and suggest that it is an important environmental factor to interpret and assess plant assemblage responses to global change.     

Terrestrial C:N stoichiometry in response to elevated CO2 and N addition: a synthesis of two meta-analyses

Both elevated atmospheric carbon dioxide (CO₂) and nitrogen (N) deposition may induce changes in C:N ratios in plant tissues and mineral soil. However, the potential mechanisms driving the stoichiometric shifts remain elusive. In this study, we examined the responses of C:N ratios in both plant tissues and mineral soil to elevated CO₂ and N deposition using data extracted from 140 peer-reviewed publications. Our results indicated that C:N ratios in both plant tissues and mineral soil exhibited consistent increases under elevated CO₂ regimes whereas decreases in C:N ratios were observed in response to experimental N addition. Moreover, soil C:N ratio was less sensitive than plant C:N ratio to both global change scenarios. Our results also showed that the responses of stoichiometric ratios were highly variable among different studies. The changes in C:N ratio did not exhibit strong correlations with C dynamics but were negatively associated with corresponding changes in N content. These results suggest that N dynamics drive stoichiometric shifts in both plant tissues and mineral soil under both elevated CO₂ and N deposition scenarios.     

Nitrogen assimilation and transpiration: key processes conditioning responsiveness of wheat to elevated [CO2] and temperature

Although climate scenarios have predicted an increase in [CO₂] and temperature conditions, to date few experiments have focused on the interaction of [CO₂] and temperature effects in wheat development. Recent evidence suggests that photosynthetic acclimation is linked to the photorespiration and N assimilation inhibition of plants exposed to elevated CO₂. The main goal of this study was to analyze the effect of interacting [CO₂] and temperature on leaf photorespiration, C/N metabolism and N transport in wheat plants exposed to elevated [CO₂] and temperature conditions. For this purpose, wheat plants were exposed to elevated [CO₂] (400 vs 700 µmol mol^?1) and temperature (ambient vs ambient + 4°C) in CO₂ gradient greenhouses during the entire life cycle. Although at the agronomic level, elevated temperature had no effect on plant biomass, physiological analyses revealed that combined elevated [CO₂] and temperature negatively affected photosynthetic performance. The limited energy levels resulting from the reduced respiratory and photorespiration rates of such plants were apparently inadequate to sustain nitrate reductase activity. Inhibited N assimilation was associated with a strong reduction in amino acid content, conditioned leaf soluble protein content and constrained leaf N status. Therefore, the plant response to elevated [CO₂] and elevated temperature resulted in photosynthetic acclimation. The reduction in transpiration rates induced limitations in nutrient transport in leaves of plants exposed to elevated [CO₂] and temperature, led to mineral depletion and therefore contributed to the inhibition of photosynthetic activity.     

Metabolite analysis of the effects of elevated CO2 and nitrogen fertilization on the association between tall fescue (Schedonorus arundinaceus) and its fungal symbiont Neotyphodium coenophialum

Atmospheric CO₂ is expected to increase to between 550 ppm and 1000 ppm in the next century. CO₂‐induced changes in plant physiology can have ecosystem‐wide implications and may alter plant‐plant, plant‐herbivore and plant‐symbiont interactions. We examined the effects of three concentrations of CO₂ (390, 800 and 1000 ppm) and two concentrations of nitrogen fertilizer (0.004 g N/week versus 0.2 g N/week) on the physiological response of Neotyphodium fungal endophyte‐infected and uninfected tall fescue plants. We used quantitative PCR to estimate the concentration of endophyte under altered CO₂ and N conditions. We found that elevated CO₂ increased the concentration of water‐soluble carbohydrates and decreased the concentration of plant total amino acids in plants. Fungal‐derived alkaloids decreased in response to elevated CO₂ and increased in response to nitrogen fertilization. Endophyte concentration (expressed as the number of copies of an endophyte‐specific gene per total genomic DNA) increased under elevated CO₂ and nitrogen fertilization. The correlation between endophyte concentration and alkaloid production observed at ambient conditions was not observed under elevated CO₂. These results suggest that nutrient exchange dynamics important for maintaining the symbiotic relationship between fungal endophytes and their grass hosts may be altered by changes in environmental variables such as CO₂ and nitrogen fertilization.     

Effects of twice-ambient carbon dioxide and nitrogen amendment on biomass, nutrient contents and carbon costs of Norway spruce seedlings as influenced by mycorrhization with Piloderma croceum and Tomentellopsis submollis

Elevated tropospheric CO₂ concentrations may increase plant carbon fixation. In ectomycorrhizal trees, a considerable portion of the synthesized carbohydrates can be used to support the mutualistic fungal root partner which in turn can benefit the tree by increased nutrient supply. In this study, Norway spruce seedlings were inoculated with either Piloderma croceum (medium distance “fringe” exploration type) or Tomentellopsis submollis (medium distance “smooth” exploration type). We studied the impact of either species regarding fungal biomass production, seedling biomass, nutrient status and nutrient use efficiency in rhizotrons under ambient and twice-ambient CO₂ concentrations. A subset was amended with ammonium nitrate to prevent nitrogen imbalances expected under growth promotion by elevated CO₂. The two fungal species exhibited considerably different influences on growth, biomass allocation as well as nutrient uptake of spruce seedlings. P. croceum increased nutrient supply and promoted plant growth more strongly than T. submollis despite considerably higher carbon costs. In contrast, seedlings with T. submollis showed higher nutrient use efficiency, i.e. produced plant biomass per received unit of nutrient, particularly for P, K and Mg, thereby promoting shoot growth and reducing the root/shoot ratio. Under the given low soil nutrient availability, P. croceum proved to be a more favourable fungal partner for seedling development than T. submollis. Additionally, plant internal allocation of nutrients was differently influenced by the two ECM fungal species, particularly evident for P in shoots and for Ca in roots. Despite slightly increased ECM length and biomass production, neither of the two species had increased its capacity of nutrient uptake in proportion to the rise of CO₂. This lead to imbalances in nutritional status with reduced nutrient concentrations, particularly in seedlings with P. croceum. The beneficial effect of P. croceum thus diminished, although the nutrient status of its host plants was still above that of plants with T. submollis. We conclude that the imbalances of nutrient status in response to elevated CO₂ at early stages of plant development are likely to prove particularly severe at nutrient-poor soils as the increased growth of ECM cannot cover the enhanced nutrient demand. Hyphal length and biomass per unit of ectomycorrhizal length as determined for the first time for P. croceum amounted to 6.9 m cm⁻¹ and 6.0 μg cm⁻¹, respectively, across all treatments.     

Linking Annual N2O Emission in Organic Soils to Mineral Nitrogen Input as Estimated by Heterotrophic Respiration and Soil C/N Ratio

Organic soils are an important source of N₂O, but global estimates of these fluxes remain uncertain because measurements are sparse. We tested the hypothesis that N₂O fluxes can be predicted from estimates of mineral nitrogen input, calculated from readily-available measurements of CO₂ flux and soil C/N ratio. From studies of organic soils throughout the world, we compiled a data set of annual CO₂ and N₂O fluxes which were measured concurrently. The input of soil mineral nitrogen in these studies was estimated from applied fertilizer nitrogen and organic nitrogen mineralization. The latter was calculated by dividing the rate of soil heterotrophic respiration by soil C/N ratio. This index of mineral nitrogen input explained up to 69% of the overall variability of N₂O fluxes, whereas CO₂ flux or soil C/N ratio alone explained only 49% and 36% of the variability, respectively. Including water table level in the model, along with mineral nitrogen input, further improved the model with the explanatory proportion of variability in N₂O flux increasing to 75%. Unlike grassland or cropland soils, forest soils were evidently nitrogen-limited, so water table level had no significant effect on N₂O flux. Our proposed approach, which uses the product of soil-derived CO₂ flux and the inverse of soil C/N ratio as a proxy for nitrogen mineralization, shows promise for estimating regional or global N₂O fluxes from organic soils, although some further enhancements may be warranted.     

The effects of CO2 and nutrient enrichment on photosynthesis and growth of Poa annua in two consecutive generations

We studied short- and long-term growth responses of Poa annua L. (Gramineae) at ambient and elevated (ambient +200???mol?mol^?1) atmospheric CO₂. In experiment 1 we compared plant growth during the early, vegetative and final, reproductive growth phases. Plant growth in elevated CO₂ was significantly enhanced during the early phase, but this was reversed in the reproductive phase. Seed mass and percentage germination were significantly reduced in elevated CO₂. Experiment 2 tested for the impact of transgenerational and nutrient effects on the response of Poa annua to elevated CO₂. Plants were grown at ambient and elevated CO₂ for one or two consecutive generations at three soil nutrient levels. Leaf photosynthesis was significantly higher at elevated CO₂, but was also affected by both soil nutrient status and plant generation. Plants grown at elevated CO₂ and under conditions of low nutrient availability showed photosynthetic acclimation after 12?weeks of growth but not after 6?weeks. First-generation growth remained unaffected by elevated CO₂, while second-generation plants produced significantly more tillers and flowers when grown in elevated CO₂ compared to ambient conditions. This effect was strongest at low nutrient availability. Average above- and belowground biomass after 12?weeks of growth was enhanced in elevated CO₂ during both generations, but more so during plant generation 2. This study demonstrates the importance of temporal/maternal effects in plant responses to elevated CO₂.     

Nutrient availability and nutrient use efficiency in plants growing in the transition zone between land and water

The transition zone between terrestrial and freshwater habitats is highly dynamic, with large variability in environmental characteristics. Here, we investigate how these characteristics influence the nutritional status and performance of plant life forms inhabiting this zone. Specifically, we hypothesised that: (i) tissue nutrient content differs among submerged, amphibious and terrestrial species, with higher content in submerged species; and (ii) PNUE gradually increases from submerged over amphibious to terrestrial species, reflecting differences in the availability of N and P relative to inorganic C across the land–water ecotone. We found that tissue nutrient content was generally higher in submerged species and C:N and C:P ratios indicated that content was limiting for growth for ca. 20% of plant individuals, particularly those belonging to amphibious and terrestrial species groups. As predicted, the PNUE increased from submerged over amphibious to terrestrial species. We suggest that this pattern reflects that amphibious and terrestrial species allocate proportionally more nutrients into processes of importance for photosynthesis at saturating CO₂ availability, i.e. enzymes involved in substrate regeneration, compared to submerged species that are acclimated to lower availability of CO₂ in the aquatic environment. Our results indicate that enhanced nutrient loading may affect relative abundance of the three species groups in the land–water ecotone of stream ecosystems. Thus, species of amphibious and terrestrial species groups are likely to benefit more from enhanced nutrient availability in terms of faster growth compared to aquatic species, and that this can be detrimental to aquatic species growing in the land–water ecotone, e.g. Ranunculus and Callitriche.     

Effects of elevated atmospheric CO2 on soil microbiota in calcareous grassland

Microbial responses to three years of CO₂ enrichment (600 μL L^–1) in the field were investigated in calcareous grassland. Microbial biomass carbon (C) and soil organic C and nitrogen (N) were not significantly influenced by elevated CO₂. Microbial C:N ratios significantly decreased under elevated CO₂ (– 15%, P = 0.01) and microbial N increased by + 18% (P = 0.04). Soil basal respiration was significantly increased on one out of 7 sampling dates (+ 14%, P = 0.03; December of the third year of treatment), whereas the metabolic quotient for CO₂ (qCO₂ = basal respiration/microbial C) did not exhibit any significant differences between CO₂ treatments. Also no responses of microbial activity and biomass were found in a complementary greenhouse study where intact grassland turfs taken from the field site were factorially treated with elevated CO₂ and phosphorus (P) fertilizer (1 g P m^–2 y^–1). Previously reported C balance calculations showed that in the ecosystem investigated growing season soil C inputs were strongly enhanced under elevated CO₂. It is hypothesized that the absence of microbial responses to these enhanced soil C fluxes originated from mineral nutrient limitations of microbial processes. Laboratory incubations showed that short-term microbial growth (one week) was strongly limited by N availability, whereas P was not limiting in this soil. The absence of large effects of elevated CO₂ on microbial activity or biomass in such nutrient-poor natural ecosystems is in marked contrast to previously published large and short-term microbial responses to CO₂ enrichment which were found in fertilized or disturbed systems. It is speculated that the absence of such responses in undisturbed natural ecosystems in which mineral nutrient cycles have equilibrated over longer periods of time is caused by mineral nutrient limitations which are ineffective in disturbed or fertilized systems and that therefore microbial responses to elevated CO₂ must be studied in natural, undisturbed systems.     

Changes in soil carbon and nitrogen properties and microbial communities in relation to growth of Pinus radiata and Nothofagus fusca trees after 6 years at ambient and elevated atmospheric CO2

Increasing atmospheric CO₂ concentration can influence the growth and chemical composition of many plant species, and thereby affect soil organic matter pools and nutrient fluxes. Here, we examine the effects of ambient (initially 362 μL L^?1) and elevated (654 μL L^?1) CO₂ in open‐top chambers on the growth after 6 years of two temperate evergreen forest species: an exotic, Pinus radiata D. Don, and a native, Nothofagus fusca (Hook. F.) Oerst. (red beech). We also examine associated effects on selected carbon (C) and nitrogen (N) properties in litter and mineral soil, and on microbial properties in rhizosphere and hyphosphere soil. The soil was a weakly developed sand that had a low initial C concentration of about 1.0 g kg^?1 at both 0–100 and 100–300 mm depths; in the N. fusca system, it was initially overlaid with about 50 mm of forest floor litter (predominantly FH material) taken from a Nothofagus forest. A slow‐release fertilizer was added during the early stages of plant growth; subsequent foliage analyses indicated that N was not limiting. After 6 years, stem diameters, foliage N concentrations and C/N ratios of both species were indistinguishable (P>0.10) in the two CO₂ treatments. Although total C contents in mineral soil at 0–100 mm depth had increased significantly (P<0.001) after 6 years growth of P. radiata, averaging 80±0.20 g m^?2 yr^?1, they were not significantly influenced by elevated CO₂. However, CO₂‐C production in litter, and CO₂‐C production, microbial C, and microbial C/N ratios in mineral soil (0–100 mm depth) under P. radiata were significantly higher under elevated than ambient CO₂. CO₂‐C production, microbial C, and numbers of bacteria (but not fungi) were also significantly higher under elevated CO₂ in hyphosphere soil, but not in rhizosphere soil. Under N. fusca, some incorporation of the overlaid litter into the mineral soil had probably occurred; except for CO₂‐C production and microbial C in hyphosphere soil, none of the biochemical properties or microbial counts increased significantly under elevated CO₂. Net mineral‐N production, and generally the potential utilization of different substrates by microbial communities, were not significantly influenced by elevated CO₂ under either tree species. Physiological profiles of the microbial communities did, however, differ significantly between rhizosphere and hyphosphere samples and between samples under P. radiata and N. fusca. Overall, results support the concept that a major effect on soil properties after prolonged exposure of trees to elevated CO₂ is an increase in the amounts, and mineralization rate, of labile organic components.     

Mineral Elements of Subtropical Tree Seedlings in Response to Elevated Carbon Dioxide and Nitrogen Addition

Mineral elements in plants have been strongly affected by increased atmospheric carbon dioxide (CO₂) concentrations and nitrogen (N) deposition due to human activities. However, such understanding is largely limited to N and phosphorus in grassland. Using open-top chambers, we examined the concentrations of potassium (K), calcium (Ca), magnesium (Mg), aluminum (Al), copper (Cu) and manganese (Mn) in the leaves and roots of the seedlings of five subtropical tree species in response to elevated CO₂ (ca. 700 μmol CO₂ mol^-1) and N addition (100 kg N ha^-1 yr^-1) from 2005 to 2009. These mineral elements in the roots responded more strongly to elevated CO₂ and N addition than those in the leaves. Elevated CO₂ did not consistently decrease the concentrations of plant mineral elements, with increases in K, Al, Cu and Mn in some tree species. N addition decreased K and had no influence on Cu in the five tree species. Given the shifts in plant mineral elements, Schima superba and Castanopsis hystrix were less responsive to elevated CO₂ and N addition alone, respectively. Our results indicate that plant stoichiometry would be altered by increasing CO₂ and N deposition, and K would likely become a limiting nutrient under increasing N deposition in subtropics.     

Genotypic variation for condensed tannin production in trembling aspen (POPULUS TREMULOIDES,salicaceae) under elevated CO2 and in high- and low-fertility soil

The carbon/nutrient balance hypothesis suggests that leaf carbon to nitrogen ratios influence the synthesis of secondary compounds such as condensed tannins. We studied the effects of rising atmospheric carbon dioxide on carbon to nitrogen ratios and tannin production. Six genotypes of Populus tremuloides were grown under elevated and ambient CO₂ partial pressure and high- and low-fertility soil in field open-top chambers in northern lower Michigan, USA. During the second year of exposure, leaves were harvested three times (June, August, and September) and analyzed for condensed tannin concentration. The carbon/nutrient balance hypothesis was supported overall, with significantly greater leaf tannin concentration at high CO₂ and low soil fertility compared to ambient CO₂ and high soil fertility. However, some genotypes increased tannin concentration at elevated compared to ambient CO₂, while others showed no CO₂ response. Performance of lepidopteran leaf miner (Phyllonorycter tremuloidiella) larvae feeding on these plants varied across genotypes, CO₂, and fertility treatments. These results suggest that with rising atmospheric CO₂, plant secondary compound production may vary within species. This could have consequences for plant–herbivore and plant–microbe interactions and for the evolutionary response of this species to global climate change.