Warming prevents the elevated CO2-induced reduction in available soil nitrogen in a temperate, perennial grassland

Rising atmospheric carbon dioxide concentration ([CO₂]) has the potential to stimulate ecosystem productivity and sink strength, reducing the effects of carbon (C) emissions on climate. In terrestrial ecosystems, increasing [CO₂] can reduce soil nitrogen (N) availability to plants, preventing the stimulation of ecosystem C assimilation; a process known as progressive N limitation. Using ion exchange membranes to assess the availability of dissolved organic N, ammonium and nitrate, we found that CO₂ enrichment in an Australian, temperate, perennial grassland did not increase plant productivity, but did reduce soil N availability, mostly by reducing nitrate availability. Importantly, the addition of 2 °C warming prevented this effect while warming without CO₂ enrichment did not significantly affect N availability. These findings indicate that warming could play an important role in the impact of [CO₂] on ecosystem N cycling, potentially overturning CO₂‐induced effects in some ecosystems.     

Warming and Nitrogen Addition Alter Photosynthetic Pigments,Sugars and Nutrients in a Temperate Meadow Ecosystem

Global warming and nitrogen (N) deposition have an important influence on terrestrial ecosystems; however, the influence of warming and N deposition on plant photosynthetic products and nutrient cycling in plants is not well understood. We examined the effects of 3 years of warming and N addition on the plant photosynthetic products, foliar chemistry and stoichiometric ratios of two dominant species, i.e., Leymus chinensis and Phragmites communis, in a temperate meadow in northeastern China. Warming significantly increased the chlorophyll content and soluble sugars in L. chinensis but had no impact on the carotenoid and fructose contents. N addition caused a significant increase in the carotenoid and fructose contents. Warming and N addition had little impact on the photosynthetic products of P. communis. Warming caused significant decreases in the N and phosphorus (P) concentrations and significantly increased the carbon (C):P and N:P ratios of L. chinensis, but not the C concentration or the C:N ratio. N addition significantly increased the N concentration, C:P and N:P ratios, but significantly reduced the C:N ratio of L. chinensis. Warming significantly increased P. communis C and P concentrations, and the C:N and C:P ratios, whereas N addition increased the C, N and P concentrations but had no impact on the stoichiometric variables. This study suggests that both warming and N addition have direct impacts on plant photosynthates and elemental stoichiometry, which may play a vital role in plant-mediated biogeochemical cycling in temperate meadow ecosystems.     

Microbial decomposition at elevated CO2 levels: effect of litter quality

The decomposition of senesced plant litter represents an important intermediate step in the cycling of nutrients between above- and below-ground systems. The rate of decomposition of plant litter is sensitive to fluctuations in a number of parameters, including environmental conditions, and particularly to changes in the quality of the litter. Increased C: N ratios of litter are thought to be one possible consequence of growth of plants under elevated [CO₂]. This response is likely to reduce the rate of decomposition of the litter. Evidence from the growth of plants in both pot and field studies suggests that growth of C3 plants in elevated atmospheric [CO₂] (600–700 μmol mol^–1) may lead to a significant increase in either/both the C: N and the lignin: N ratios of litter. Short-term decomposition of litter from plants showing this response in elevated [CO₂] has confirmed that decomposition occurs at a significantly lower rate. The limited studies of both the response of C4 plants to elevated [CO₂] and the subsequent degradability of the senescent litter suggest that no differences in litter quality or degradability occur. In terms of litter quality the response of plants therefore appears to be dependent upon photosynthetic type; the C:N and lignin:N ratios of litter from C3 plants exposed to elevated [CO₂] are increased, leading to lower degradation rates, while the nutrient ratios and degradation rates of litter from C4 plants grown in elevated [CO₂] remain unchanged. To date, very few ecosystem studies of decomposition have been carried out. Further work is required at the ecosystem level to determine whether the effects observed in laboratory, pot and field studies are also observed in long-term, complex ecosystem studies. Clearly if these results are repeated at the ecosystem level then significant changes in the cycling of C and N in important terrestrial ecosystems may occur as a results of elevated [CO₂].     

Effects of three global change drivers on terrestrial C:N:P stoichiometry: a global synthesis

Over the last few decades, there has been an increasing number of controlled‐manipulative experiments to investigate how plants and soils might respond to global change. These experiments typically examined the effects of each of three global change drivers [i.e., nitrogen (N) deposition, warming, and elevated CO₂] on primary productivity and on the biogeochemistry of carbon (C), N, and phosphorus (P) across different terrestrial ecosystems. Here, we capitalize on this large amount of information by performing a comprehensive meta‐analysis (>2000 case studies worldwide) to address how C:N:P stoichiometry of plants, soils, and soil microbial biomass might respond to individual vs. combined effects of the three global change drivers. Our results show that (i) individual effects of N addition and elevated CO₂ on C:N:P stoichiometry are stronger than warming, (ii) combined effects of pairs of global change drivers (e.g., N addition + elevated CO₂, warming + elevated CO₂) on C:N:P stoichiometry were generally weaker than the individual effects of each of these drivers, (iii) additive interactions (i.e., when combined effects are equal to or not significantly different from the sum of individual effects) were more common than synergistic or antagonistic interactions, (iv) C:N:P stoichiometry of soil and soil microbial biomass shows high homeostasis under global change manipulations, and (v) C:N:P responses to global change are strongly affected by ecosystem type, local climate, and experimental conditions. Our study is one of the first to compare individual vs. combined effects of the three global change drivers on terrestrial C:N:P ratios using a large set of data. To further improve our understanding of how ecosystems might respond to future global change, long‐term ecosystem‐scale studies testing multifactor effects on plants and soils are urgently required across different world regions.     

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.     

Anthropogenic global shifts in biospheric N and P concentrations and ratios and their impacts on biodiversity,ecosystem productivity,food security,and human health

The availability of carbon (C) from high levels of atmospheric carbon dioxide (CO₂) and anthropogenic release of nitrogen (N) is increasing, but these increases are not paralleled by increases in levels of phosphorus (P). The current unstoppable changes in the stoichiometries of C and N relative to P have no historical precedent. We describe changes in P and N fluxes over the last five decades that have led to asymmetrical increases in P and N inputs to the biosphere. We identified widespread and rapid changes in N:P ratios in air, soil, water, and organisms and important consequences to the structure, function, and biodiversity of ecosystems. A mass‐balance approach found that the combined limited availability of P and N was likely to reduce C storage by natural ecosystems during the remainder of the 21st Century, and projected crop yields of the Millennium Ecosystem Assessment indicated an increase in nutrient deficiency in developing regions if access to P fertilizer is limited. Imbalances of the N:P ratio would likely negatively affect human health, food security, and global economic and geopolitical stability, with feedbacks and synergistic effects on drivers of global environmental change, such as increasing levels of CO₂, climatic warming, and increasing pollution. We summarize potential solutions for avoiding the negative impacts of global imbalances of N:P ratios on the environment, biodiversity, climate change, food security, and human health.     

Nitrogen:phosphorous supply ratio and allometry in five alpine plant species

In terrestrial ecosystems, atmospheric nitrogen (N) deposition has greatly increased N availability relative to other elements, particularly phosphorus (P). Alterations in the availability of N relative to P can affect plant growth rate and functional traits, as well as resource allocation to above‐ versus belowground biomass (M_A and M_B). Biomass allocation among individual plants is broadly size‐dependent, and this can often be described as an allometric relationship between M_A and M_B, as represented by the equation , or log M_A = logα + βlog M_B. Here, we investigated whether the scaling exponent or regression slope may be affected by the N:P supply ratio. We hypothesized that the regression slope between M_A and M_B should be steeper under a high N:P supply ratio due to P limitation, and shallower under a low N:P supply ratio due to N limitation. To test these hypotheses, we experimentally altered the levels of N, P, and the N:P supply ratio (from 1.7:1 to 135:1) provided to five alpine species representing two functional groups (grasses and composite forbs) under greenhouse conditions; we then measured the effects of these treatments on plant morphology and tissue content (SLA, leaf area, and leaf and root N/P concentrations) and on the scaling relationship between M_A and M_B. Unbalanced N:P supply ratios generally negatively affected plant biomass, leaf area, and tissue nutrient concentration in both grasses and composite forbs. High N:P ratios increased tissue N:P ratios in both functional groups, but more in the two composite forbs than in the grasses. The positive regression slopes between log M_A and log M_B exhibited by plants raised under a N:P supply ratio of 135:1 were significantly steeper than those observed under the N:P ratio of 1.7:1 and 15:1. Synthesis: Plant biomass allocation is highly plastic in response to variation in the N:P supply ratio. Studies of resource allocation of individual plants should focus on the effects of nutrient ratios as well as the availability of individual elements. The two forb species were more sensitive than grasses to unbalanced N:P supplies. To evaluate the adaptive significance of this plasticity, the effects of unbalanced N:P supply ratio on individual lifetime fitness must be measured.     

Stoichiometric response of nitrogen-fixing and non-fixing dicots to manipulations of CO2, nitrogen, and diversity

Human activities have resulted in increased nitrogen deposition and atmospheric CO₂ concentrations in the biosphere, potentially causing significant changes in many ecological processes. In addition to these ongoing perturbations of the abiotic environment, human-induced losses of biodiversity are also of major concern and may interact in important ways with biogeochemical perturbations to affect ecosystem structure and function. We have evaluated the effects of these perturbations on plant biomass stoichiometric composition (C:N:P ratios) within the framework of the BioCON experimental setup (biodiversity, CO₂, N) conducted at the Cedar Creek Natural History Area, Minnesota. Here we present data for five plant species: Solidago rigida, Achillea millefolium, Amorpha canescens, Lespedeza capitata, and Lupinus perennis. We found significantly higher C:N and C:P ratios under elevated CO₂ treatments, but species responded idiosyncratically to the treatment. Nitrogen addition decreased C:N ratios, but this response was greater in the ambient CO₂ treatments than under elevated CO₂. Higher plant species diversity generally lowered both C:N and C:P ratios. Importantly, increased diversity also led to a more modest increase in the C:N ratio with elevated CO₂ levels. In addition, legumes exhibited lower C:N and higher C:P and N:P ratios than non-legumes, highlighting the effect of physiological characteristics defining plant functional types. These data suggest that atmospheric CO₂ levels, N availability, and plant species diversity interact to affect both aboveground and belowground processes by altering plant elemental composition.     

ELEVATED ATMOSPHERIC CARBON DIOXIDE INCREASES ORGANIC CARBON FIXATION BY EMILIANIA HUXLEYI (HAPTOPHYTA), UNDER NUTRIENT‐LIMITED HIGH‐LIGHT CONDITIONS1

Phytoplankton play a key role in determining the partitioning of CO₂ between the atmosphere and the ocean on seasonal, interannual, and millennial time scales. The magnitude of biological draw‐down of atmospheric CO₂ and C storage in the oceans is affected by concurrent changes in other environmental factors, like nutrient supply. Furthermore, variations in carbon‐to‐nitrogen (C:N) and carbon‐to‐phosphorus (C:P) assimilation ratios modify the oceanic CO₂ storage capacity. Here we show that increased atmospheric CO₂ concentration enhances CO₂ fixation into organic matter by a noncalcifying strain of Emiliania huxleyi (Lohmann) Hay & Mohler only under certain conditions, namely high light and nutrient limitation. Enhanced organic matter production was accompanied by marked deviations of the C:N:P ratio from the canonical stoichiometry of marine particulate matter of 106:16:1 (C:N:P) known as the Redfield ratio. Increased cell organic carbon content, C:N, and C:P were observed at high light when growth was either nitrogen or phosphorus limited. Elevated CO₂ led to further increases in the particulate C:N and C:P ratios. Enhanced CO₂ uptake by phytoplankton such as E. huxleyi, in response to elevated atmospheric CO₂, could increase carbon storage in the nitrogen‐limited regions of the oceans and thus act as a negative feedback on rising atmospheric CO₂ levels.     

Decomposition of litter produced under elevated CO2: Dependence on plant species and nutrient supply

We investigated the effect of CO₂ concentration and soilnutrient availability during growth on the subsequent decomposition andnitrogen (N) release from litter of four annual grasses that differ inresource requirements and native habitat. Vulpia microstachys isa native grass found on California serpentine soils, whereas Avenafatua, Bromus hordaceus, and Lolium multiflorum areintroduced grasses restricted to more fertile sandstone soils (Hobbs & Mooney 1991). Growth in elevated CO₂ altered litter C:N ratio,decomposition, and N release, but the direction and magnitude of thechanges differed among plant species and nutrient treatments. ElevatedCO₂ had relatively modest effects on C:N ratio of litter,increasing this ratio in Lolium roots (and shoots at high nutrients),but decreasing C:N ratio in Avena shoots. Growth of plants underelevated CO₂ decreased the decomposition rate of Vulpialitter, but increased decomposition of Avena litter from the high-nutrient treatment. The impact of elevated CO₂ on N loss fromlitter also differed among species, with Vulpia litter from high-CO₂ plants releasing N more slowly than ambient-CO₂litter, whereas growth under elevated CO₂ caused increased Nloss from Avena litter. CO₂ effects on N release in Lolium and Bromus depended on the nutrient regime in whichplants were grown. There was no overall relationship between litter C:Nratio and decomposition rate or N release across species and treatments.Based on our study and the literature, we conclude that the effects ofelevated CO₂ on decomposition and N release from litter arehighly species-specific. These results do not support the hypothesis thatCO₂ effects on litter quality consistently lead to decreasednutrient availability in nutrient-limited ecosystems exposed to elevatedCO₂.     

Increases of Soil C, N, and P Pools Along an Acacia Tree Density Gradient and Their Effects on Trees and Grasses

Nitrogen (N) fixing trees including many species of Acacia are an important though variable component of savanna ecosystems. It is known that these trees enrich the soil with carbon (C) and N, but their effect on the combined C:N:P stoichiometry in soil is less well understood. Theory suggests that they might reduce available phosphorus (P), creating a shift from more N-limited conditions in grass-dominated to more P-limited conditions in tree-dominated sites, which in turn could feed back negatively on the trees’ capacity to fix N. We studied the effects of Acacia zanzibarica tree density upon soil and foliar N:P stoichiometry, and the N₂-fixation rates of trees and leguminous herbs in a humid Tanzanian savanna. Foliar N:P ratios and N₂-fixation rates of trees remained constant across the density gradient, whereas soil C, N and organic P pools increased. In contrast, the N:P ratio of grasses increased and N₂-fixation rates of leguminous herbs decreased with increasing tree density, indicating a shift towards more P-limited conditions for the understory vegetation. These contrasting responses suggest that trees and grasses have access to different sources of N and P, with trees being able to access P from deeper soil layers and perhaps also utilizing organic forms more efficiently.     

Soil C:N:P stoichiometry responds to vegetation change from grassland to woodland

Woody encroachment has been a major land cover change in dryland ecosystems during the past century. While numerous studies have demonstrated strong effects of woody encroachment on soil carbon (C), nitrogen (N), and phosphorus (P) storage, far less is known about the plasticity of soil C:N:P stoichiometry in response to woody encroachment. We assessed landscape-scale patterns of spatial heterogeneity in soil C:N:P ratios throughout a 1.2 m soil profile in a region where grassland is being replaced by a diverse assemblage of subtropical woody plants dominated by Prosopis glandulosa, an N₂-fixing tree. Woody species had leaf and fine root C:N:P ratios significantly different from grasses. Variation in soil C:N ratios in both horizontal and vertical planes was remarkably smaller than that of soil N:P and C:P ratios. Spatial patterns of soil C:N ratio throughout the profile were not strongly related to vegetation cover. In contrast, spatial patterns of soil N:P and C:P ratios displayed a strong resemblance to that of vegetation cover throughout the soil profile. Within the uppermost soil layer (0–5 cm), soil N:P and C:P ratios were higher underneath woody patches while lower within the grassland; however, this pattern was reversed in subsurface soils (15–120 cm). These results indicate a complex response of soil C:N:P stoichiometry to vegetation change, which could have important implications for understanding C, N, and P interactions and nutrient limitations in dryland ecosystems.     

Laboratory incubations reveal potential responses of soil nitrogen cycling to changes in soil C and N availability in Mojave Desert soils exposed to elevated atmospheric CO2

Elevated atmospheric carbon dioxide (CO₂) has the potential to alter soil carbon (C) and nitrogen (N) cycling in arid ecosystems through changes in net primary productivity. However, an associated feedback exists because any sustained increases in plant productivity will depend upon the continued availability of soil N. We took soils from under the canopies of major shrubs, grasses, and plant interspaces in a Mojave Desert ecosystem exposed to elevated atmospheric CO₂ and incubated them in the laboratory with amendments of labile C and N to determine if elevated CO₂ altered the mechanistic controls of soil C and N on microbial N cycling. Net ammonification increased under shrubs exposed to elevated CO₂, while net nitrification decreased. Elevated CO₂ treatments exhibited greater fluxes of N₂O–N under Lycium spp., but not other microsites. The proportion of microbial/extractable organic N increased under shrubs exposed to elevated CO₂. Heterotrophic N₂‐fixation and C mineralization increased with C addition, while denitrification enzyme activity and N₂O–N fluxes increased when C and N were added in combination. Laboratory results demonstrated the potential for elevated CO₂ to affect soil N cycling under shrubs and supports the hypothesis that energy limited microbes may increase net inorganic N cycling rates as the amount of soil‐available C increases under elevated CO₂. The effect of CO₂ enrichment on N‐cycling processes is mediated by its effect on the plants, particularly shrubs. The potential for elevated atmospheric CO₂ to lead to accumulation of NH₄⁺ under shrubs and the subsequent volatilization of NH₃ may result in greater losses of N from this system, leading to changes in the form and amount of plant‐available inorganic N. This introduces the potential for a negative feedback mechanism that could act to constrain the degree to which plants can increase productivity in the face of elevated atmospheric CO₂.     

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

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

Nitrogen cycling in microcosms of yellow birch exposed to elevated CO2: simultaneous positive and negative below-ground feedbacks

This study investigated simultaneous plant and soil feedbacks on growth enhancement with elevated [CO₂] within microcosms of yellow birch (Betula alleghaniensis Britt.) in the second year of growth. Understanding the integrated responses of model ecosystems may provide key insight into the potential net nutrient feedbacks on [CO₂] growth enhancements in temperate forests. We measured the net biomass production, C:N ratios, root architecture, and mycorrhizal responses of yellow birch, in situ rates gross nitrogen mineralization and the partitioning of available NH₄⁺ between yellow birch and soil microbes. Elevated atmospheric [CO₂] resulted in significant alterations in the cycling of N within the microcosms. Plant C/N ratios were significantly increased, gross mineralization and NH₄⁺ consumption rates were decreased, and relative microbial uptake of NH₄⁺ was increased, representing a suite of N cycling negative feedbacks on N availability. However, increased C/N ratios may also be a mechanism which allows plants to maintain higher growth with a constant or reduced N supply. Total plant N content was increased with elevated [CO₂], suggesting that yellow birch had successfully increased their ability to acquire nutrients during the first year of growth. However, plant uptake rates of NH₄⁺ had decreased in the second year. This discrepancy implies that, in this study, nitrogen uptake showed a trend through ontogeny of decreasing enhancement under elevated [CO₂]. The reduced N mineralization and relatively increased N immobilization are a potential feedback which may drive this ontogenetic trend. This study has demonstrated the importance of using an integrated approach to exploring potential nutrient-cycling feedbacks in elevated [CO₂].     

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 availability mediates soil carbon cycling response to climate warming: A meta-analysis

Global climate warming may induce a positive feedback through increasing soil carbon (C) release to the atmosphere. Although warming can affect both C input to and output from soil, direct and convincing evidence illustrating that warming induces a net change in soil C is still lacking. We synthesized the results from field warming experiments at 165 sites across the globe and found that climate warming had no significant effect on soil C stock. On average, warming significantly increased root biomass and soil respiration, but warming effects on root biomass and soil respiration strongly depended on soil nitrogen (N) availability. Under high N availability (soil C:N ratio < 15), warming had no significant effect on root biomass, but promoted the coupling between effect sizes of root biomass and soil C stock. Under relative N limitation (soil C:N ratio > 15), warming significantly enhanced root biomass. However, the enhancement of root biomass did not induce a corresponding C accumulation in soil, possibly because warming promoted microbial CO₂ release that offset the increased root C input. Also, reactive N input alleviated warming-induced C loss from soil, but elevated atmospheric CO₂ or precipitation increase/reduction did not. Together, our findings indicate that the relative availability of soil C to N (i.e., soil C:N ratio) critically mediates warming effects on soil C dynamics, suggesting that its incorporation into C-climate models may improve the prediction of soil C cycling under future global warming scenarios.     

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.     

Relationships between C3 Plant Foliar Carbon Isotope Composition and Element Contents of Grassland Species at High Altitudes on the Qinghai-Tibet Plateau,China

Relationships of foliar carbon isotope composition (δ¹³C) with foliar C, N, P, K, Ca, Mg contents and their ratios of 219 C₃ species leaf samples, obtained in August in 2004 to 2007 from 82 high altitude grassland sites on the Qinghai-Tibet Plateau China, were examined. This was done with reference to the proposition that foliar δ¹³C increases with altitude and separately for the life-form groups of graminoids, forbs and shrubs and for the genera Stipa and Kobresia. For all samples, foliar δ¹³C was negatively related to foliar K, P and ∑_{K+ Ca+ Mg}, and positively correlated to foliar C, C/N and C/P. The significance of these correlations differed for the taxonomic and life-form groups. Lack of a relationship of foliar δ¹³C with foliar N was inconsistent with the majority of studies that have shown foliar δ¹³C to be positively related to foliar N due to a decrease of C_i/C_a (the ratio between intercellular and atmospheric concentration of CO₂) and explained as a result of greater photosynthetic capacity at higher foliar N concentration. However this inconsistency relates to other high altitude studies that have found that photosynthetic capacity remains constant as foliar N increases. After accounting for the altitudinal relationship with foliar δ¹³C, of the elements only the K effect was significant and was most strongly expressed for Kobresia. It is concluded that factors critical to plant survival and growth at very high altitudes, such as low atmospheric pressure and low temperatures, may preclude expression of relationships between foliar δ¹³C and foliar elements that have been observed at lower altitudes.     

Individual and interactive effects of warming and nitrogen supply on CO2 fluxes and carbon allocation in subarctic grassland

Climate warming has been suggested to impact high latitude grasslands severely, potentially causing considerable carbon (C) losses from soil. Warming can also stimulate nitrogen (N) turnover, but it is largely unclear whether and how altered N availability impacts belowground C dynamics. Even less is known about the individual and interactive effects of warming and N availability on the fate of recently photosynthesized C in soil. On a 10-year geothermal warming gradient in Iceland, we studied the effects of soil warming and N addition on CO₂ fluxes and the fate of recently photosynthesized C through CO₂ flux measurements and a ¹³CO₂ pulse-labeling experiment. Under warming, ecosystem respiration exceeded maximum gross primary productivity, causing increased net CO₂ emissions. N addition treatments revealed that, surprisingly, the plants in the warmed soil were N limited, which constrained primary productivity and decreased recently assimilated C in shoots and roots. In soil, microbes were increasingly C limited under warming and increased microbial uptake of recent C. Soil respiration was increased by warming and was fueled by increased belowground inputs and turnover of recently photosynthesized C. Our findings suggest that a decade of warming seemed to have induced a N limitation in plants and a C limitation by soil microbes. This caused a decrease in net ecosystem CO₂ uptake and accelerated the respiratory release of photosynthesized C, which decreased the C sequestration potential of the grassland. Our study highlights the importance of belowground C allocation and C-N interactions in the C dynamics of subarctic ecosystems in a warmer world.