Responses of fungal root colonization,plant cover and leaf nutrients to long‐term exposure to elevated atmospheric CO2 and warming in a subarctic birch forest understory

Responses of the mycorrhizal fungal community in terrestrial ecosystems to global change factors are not well understood. However, virtually all land plants form symbiotic associations with mycorrhizal fungi, with approximately 20% of the plants' net primary production transported down to the fungal symbionts. In this study, we investigated how ericoid mycorrhiza (ErM), fine endophytes (FE) and dark septate endophytes (DSE) in roots responded to elevated atmospheric CO₂ concentrations and warming in the dwarf shrub understory of a birch forest in the subarctic region of northern Sweden. To place the belowground results into an ecosystem context we also investigated how plant cover and nutrient concentrations in leaves responded to elevated atmospheric CO₂ concentrations and warming. The ErM colonization in ericaceous dwarf shrubs increased under elevated atmospheric CO₂ concentrations, but did not respond to warming following 6 years of treatment. This suggests that the higher ErM colonization under elevated CO₂ might be due to increased transport of carbon belowground to acquire limiting resources such as N, which was diluted in leaves of ericaceous plants under enhanced CO₂. The elevated CO₂ did not affect total plant cover but the plant cover was increased under warming, which might be due to increased N availability in soil. FE colonization in grass roots decreased under enhanced CO₂ and under warming, which might be due to increased root growth, to which the FE fungi could not keep up, resulting in proportionally lower colonization. However, no responses in aboveground cover of Deschampsia flexuosa were seen. DSE hyphal colonization in grass roots significantly increased under warmer conditions, but did not respond to elevated CO₂. This complex set of responses by mycorrhizal and other root‐associated fungi to global change factors of all the fungal types studied could have broad implications for plant community structure and biogeochemistry of subarctic ecosystems.     

Do elevations in temperature,CO2, and nutrient availability modify belowground carbon gain and root morphology in artificially defoliated silver birch seedlings?

Climate warming increases the risk of insect defoliation in boreal forests. Losses in photosynthetically active surfaces cause reduction in net primary productivity and often compromise carbon reserves of trees. The concurrent effects of climate change and removal of foliage on root growth responses and carbohydrate dynamics are poorly understood, especially in tree seedlings. We investigated if exposures to different combinations of elevated temperature, CO₂, and nutrient availability modify belowground carbon gain and root morphology in artificially defoliated 1‐year‐old silver birches (Betula pendula). We quantified nonstructural carbohydrates (insoluble starch as a storage compound; soluble sucrose, fructose, and glucose) singly and in combination in fine roots of plants under winter dormancy. Also the total mass, fine root proportion, water content, and length of roots were defined. We hypothesized that the measured properties are lower in defoliated birch seedlings that grow with ample resources than with scarce resources. On average, fertilization markedly decreased both the proportion and the carbohydrate concentrations of fine roots in all seedlings, whereas the effect of fertilization on root water content and dry mass was the opposite. However, defoliation mitigated the effect of fertilization on the root water content, as well as on the proportion of fine roots and their carbohydrate concentrations by reversing the outcomes. Elevation in temperature decreased and elevation in CO₂ increased the absolute contents of total nonstructural carbohydrates, whereas fertilization alleviated both these effects. Also the root length and mass increased by CO₂ elevation. This confirms that surplus carbon in birch tissues is used as a substrate for storage compounds and for cell wall synthesis. To conclude, our results indicate that some, but not all elements of climate change alter belowground carbon gain and root morphology in defoliated silver birch seedlings.     

Carbon cost of plant nitrogen acquisition: global carbon cycle impact from an improved plant nitrogen cycle in the Community Land Model

Plants typically expend a significant portion of their available carbon (C) on nutrient acquisition – C that could otherwise support growth. However, given that most global terrestrial biosphere models (TBMs) do not include the C cost of nutrient acquisition, these models fail to represent current and future constraints to the land C sink. Here, we integrated a plant productivity‐optimized nutrient acquisition model – the Fixation and Uptake of Nitrogen Model – into one of the most widely used TBMs, the Community Land Model. Global plant nitrogen (N) uptake is dynamically simulated in the coupled model based on the C costs of N acquisition from mycorrhizal roots, nonmycorrhizal roots, N‐fixing microbes, and retranslocation (from senescing leaves). We find that at the global scale, plants spend 2.4 Pg C yr^?1 to acquire 1.0 Pg N yr^?1, and that the C cost of N acquisition leads to a downregulation of global net primary production (NPP) by 13%. Mycorrhizal uptake represented the dominant pathway by which N is acquired, accounting for ~66% of the N uptake by plants. Notably, roots associating with arbuscular mycorrhizal (AM) fungi – generally considered for their role in phosphorus (P) acquisition – are estimated to be the primary source of global plant N uptake owing to the dominance of AM‐associated plants in mid‐ and low‐latitude biomes. Overall, our coupled model improves the representations of NPP downregulation globally and generates spatially explicit patterns of belowground C allocation, soil N uptake, and N retranslocation at the global scale. Such model improvements are critical for predicting how plant responses to altered N availability (owing to N deposition, rising atmospheric CO₂, and warming temperatures) may impact the land C sink.     

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.     

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     

No cumulative effect of 10 years of elevated [CO2] on perennial plant biomass components in the Mojave Desert

Elevated atmospheric CO₂ concentrations ([CO₂]) generally increase primary production of terrestrial ecosystems. Production responses to elevated [CO₂] may be particularly large in deserts, but information on their long‐term response is unknown. We evaluated the cumulative effects of elevated [CO₂] on primary production at the Nevada Desert FACE (free‐air carbon dioxide enrichment) Facility. Aboveground and belowground perennial plant biomass was harvested in an intact Mojave Desert ecosystem at the end of a 10‐year elevated [CO₂] experiment. We measured community standing biomass, biomass allocation, canopy cover, leaf area index (LAI), carbon and nitrogen content, and isotopic composition of plant tissues for five to eight dominant species. We provide the first long‐term results of elevated [CO₂] on biomass components of a desert ecosystem and offer information on understudied Mojave Desert species. In contrast to initial expectations, 10 years of elevated [CO₂] had no significant effect on standing biomass, biomass allocation, canopy cover, and C : N ratios of above‐ and belowground components. However, elevated [CO₂] increased short‐term responses, including leaf water‐use efficiency (WUE) as measured by carbon isotope discrimination and increased plot‐level LAI. Standing biomass, biomass allocation, canopy cover, and C : N ratios of above‐ and belowground pools significantly differed among dominant species, but responses to elevated [CO₂] did not vary among species, photosynthetic pathway (C₃ vs. C₄), or growth form (drought‐deciduous shrub vs. evergreen shrub vs. grass). Thus, even though previous and current results occasionally show increased leaf‐level photosynthetic rates, WUE, LAI, and plant growth under elevated [CO₂] during the 10‐year experiment, most responses were in wet years and did not lead to sustained increases in community biomass. We presume that the lack of sustained biomass responses to elevated [CO₂] is explained by inter‐annual differences in water availability. Therefore, the high frequency of low precipitation years may constrain cumulative biomass responses to elevated [CO₂] in desert environments.     

The C:N:P stoichiometry of organisms and ecosystems in a changing world: A review and perspectives

This study examined the literature in ISI Web of Science to identify the effects that the main drivers of global change have on the nutrient concentrations and C:N:P stoichiometry of organisms and ecosystems, and examined their relationship to changes in ecosystem structure and function. We have conducted a meta-analysis by comparing C:N:P ratios of plants and soils subjected to elevated [CO₂] with those subjected to ambient [CO₂]. A second meta-analysis compared the C:N:P ratios of plants and soils that received supplemental N to simulate N deposition and those that did not receive supplemental N. On average, an experimental increase in atmospheric [CO₂] increased the foliar C:N ratios of C3 grasses, forbs, and woody plants by 22%, but the foliar ratios of C4 grasses were unaffected. This trend may be enhanced in semi-arid areas by the increase in droughts that have been projected for the coming decades which can increase leaf C:N ratios. The available studies show an average 38% increase in foliar C:P ratios in C3 plants in response to elevated atmospheric [CO₂], but no significant effects were observed in C4 grasses. Furthermore, studies that examine the effects of elevated atmospheric [CO₂] on N:P ratio (on a mass basis) are warranted since its response remains elusive. N deposition increases the N:P ratio in the plants of terrestrial and freshwater ecosystems, and decreases plants and organic soil C:N ratio (25% on average for C3 plants), reducing soil and water N₂ fixation capacity and ecosystem species diversity. In contrast, in croplands subjected to intense fertilization, mostly, animal slurries, a reduction in soil N:P ratio can occur because of the greater solubility and loss of N. In the open ocean, there are experimental observations showing an ongoing increase in P-limited areas in response to several of the factors that promote global change, including the increase in atmospheric [CO₂] which increases the demand for P, the warming effect that leads to an increase in water column stratification, and increases in the N:P ratio of atmospheric inputs. Depending on the type of plant and the climate where it grows, warming can increase, reduce, or have no effect on foliar C:N ratios. The results suggest that warming and drought can increase C:N and C:P ratios in warm-dry and temperate-dry terrestrial ecosystems, especially, when high temperatures and drought coincide. Advances in this topic are a challenge because changes in stoichiometric ratios can favour different types of species and change ecosystem composition and structure.     

Influence of carbon dioxide enrichment, ozone and nitrogen fertilization on cotton (Gossypium hirsutum L.) leaf and root composition

The objective of this study was to test whether elevated [CO₂], [O₃] and nitrogen (N) fertility altered leaf mass per area (LMPA), non‐structural carbohydrate (TNC), N, lignin (LTGA) and proanthocyanidin (PA) concentrations in cotton (Gossypium hirsutum L.) leaves and roots. Cotton was grown in 14 dm³ pots with either sufficient (0·8 g N dm ^{? 3}) or deficient (0·4 and 0·2 g N dm ^{? 3}) N fertilization, and treated in open‐top chambers with either ambient or elevated ( + 175 and + 350 μ mol mol ^{? 1}) [CO₂] in combination with either charcoal‐filtered air (CF) or non‐filtered air plus 1·5 times ambient [O₃]. At about 50 d after planting, LMPA, starch and PA concentrations in canopy leaves were as much as 51–72% higher in plants treated with elevated [CO₂] compared with plants treated with ambient [CO₂], whereas leaf N concentration was 29% lower in elevated [CO₂]‐treated plants compared with controls. None of the treatments had a major effect on LTGA concentrations on a TNC‐free mass basis. LMPA and starch levels were up to 48% lower in plants treated with elevated [O₃] and ambient [CO₂] compared with CF controls, although the elevated [O₃] effect was diminished when plants were treated concurrently with elevated [CO₂]. On a total mass basis, leaf N and PA concentrations were higher in samples treated with elevated [O₃] in ambient [CO₂], but the difference was much reduced by elevated [CO₂]. On a TNC‐free basis, however, elevated [O₃] had little effect on tissue N and PA concentrations. Fertilization treatments resulted in higher PA and lower N concentrations in tissues from the deficient N fertility treatments. The experiment showed that suppression by elevated [O₃] of LMPA and starch was largely prevented by elevated [CO₂], and that interpretation of [CO₂] and [O₃] effects should include comparisons on a TNC‐free basis. Overall, the experiment indicated that allocation to starch and PA may be related to how environmental factors affect source–sink relationships in plants, although the effects of elevated [O₃] on secondary metabolites differed in this respect.     

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

We tested a conceptual model describing the influence of elevated atmospheric CO₂ on plant production, soil microorganisms, and the cycling of C and N in the plant-soil system. Our model is based on the observation that in nutrient-poor soils, plants (C₃) grown in an elevated CO₂ atmosphere often increase production and allocation to belowground structures. We predicted that greater belowground C inputs at elevated CO₂ should elicit an increase in soil microbial biomass and increased rates of organic matter turnover and nitrogen availability. We measured photosynthesis, biomass production, and C allocation of Populus grandidentata Michx. grown in nutrient-poor soil for one field season at ambient and twice-ambient (i.e., elevated) atmospheric CO₂ concentrations. Plants were grown in a sandy subsurface soil i) at ambient CO₂ with no open top chamber, ii) at ambient CO₂ in an open top chamber, and iii) at twice-ambient CO₂ in an open top chamber. Plants were fertilized with 4.5 g N m⁻² over a 47 d period midway through the growing season. Following 152 d of growth, we quantified microbial biomass and the availabilities of C and N in rhizosphere and bulk soil. We tested for a significant CO₂ effect on plant growth and soil C and N dynamics by comparing the means of the chambered ambient and chambered elevated CO₂ treatments. Rates of photosynthesis in plants grown at elevated CO₂ were significantly greater than those measured under ambient conditions. The number of roots, root length, and root length increment were also substantially greater at elevated CO₂. Total and belowground biomass were significantly greater at elevated CO₂. Under N-limited conditions, plants allocated 50–70% of their biomass to roots. Labile C in the rhizosphere of elevated-grown plants was significantly greater than that measured in the ambient treatments; there were no significant differences between labile C pools in the bulk soil of ambient and elevated-grown plants. Microbial biomass C was significantly greater in the rhizosphere and bulk soil of plants grown at elevated CO₂ compared to that in the ambient treatment. Moreover, a short-term laboratory assay of N mineralization indicated that N availability was significantly greater in the bulk soil of the elevated-grown plants. Our results suggest that elevated atmospheric CO₂ concentrations can have a positive feedback effect on soil C and N dynamics producing greater N availability. Experiments conducted for longer periods of time will be necessary to test the potential for negative feedback due to altered leaf litter chemistry. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

Global change belowground: impacts of elevated CO2, nitrogen,and summer drought on soil food webs and biodiversity

The world's ecosystems are subjected to various anthropogenic global change agents, such as enrichment of atmospheric CO₂ concentrations, nitrogen (N) deposition, and changes in precipitation regimes. Despite the increasing appreciation that the consequences of impending global change can be better understood if varying agents are studied in concert, there is a paucity of multi‐factor long‐term studies, particularly on belowground processes. Herein, we address this gap by examining the responses of soil food webs and biodiversity to enrichment of CO₂, elevated N, and summer drought in a long‐term grassland study at Cedar Creek, Minnesota, USA (BioCON experiment). We use structural equation modeling (SEM), various abiotic and biotic explanatory variables, and data on soil microorganisms, protozoa, nematodes, and soil microarthropods to identify the impacts of multiple global change effects on drivers belowground. We found that long‐term (13‐year) changes in CO₂ and N availability resulted in modest alterations of soil biotic food webs and biodiversity via several mechanisms, encompassing soil water availability, plant productivity, and – most importantly – changes in rhizodeposition. Four years of manipulation of summer drought exerted surprisingly minor effects, only detrimentally affecting belowground herbivores and ciliate protists at elevated N. Elevated CO₂ increased microbial biomass and the density of ciliates, microarthropod detritivores, and gamasid mites, most likely by fueling soil food webs with labile C. Moreover, beneficial bottom‐up effects of elevated CO₂ compensated for detrimental elevated N effects on soil microarthropod taxa richness. In contrast, nematode taxa richness was lowest at elevated CO₂ and elevated N. Thus, enrichment of atmospheric CO₂ concentrations and N deposition may result in taxonomically and functionally altered, potentially simplified, soil communities. Detrimental effects of N deposition on soil biodiversity underscore recent reports on plant community simplification. This is of particular concern, as soils house a considerable fraction of global biodiversity and ecosystem functions.     

Interactive Effects of Time, CO2, N, and Diversity on Total Belowground Carbon Allocation and Ecosystem Carbon Storage in a Grassland Community

Predicting if ecosystems will mitigate or exacerbate rising CO₂ requires understanding how elevated CO₂ will interact with coincident changes in diversity and nitrogen (N) availability to affect ecosystem carbon (C) storage. Yet achieving such understanding has been hampered by the difficulty of quantifying belowground C pools and fluxes. Thus, we used mass balance calculations to quantify the effects of diversity, CO₂, and N on both the total amount of C allocated belowground by plants (total belowground C allocation, TBCA) and ecosystem C storage in a periodically burned, 8-year Minnesota grassland biodiversity, CO₂, and N experiment (BioCON). Annual TBCA increased in response to elevated CO₂, enriched N, and increasing diversity. TBCA was positively related to standing root biomass. After removing the influence of root biomass, the effect of elevated CO₂ remained positive, suggesting additional drivers of TBCA apart from those that maintain high root biomass. Removing root biomass effects resulted in the effects of N and diversity becoming neutral or negative (depending on year), suggesting that the positive effects of diversity and N on TBCA were related to treatment-driven differences in root biomass. Greater litter production in high diversity, elevated CO₂, and enhanced N treatments increased annual ecosystem C loss in fire years and C gain in non-fire years, resulting in overall neutral C storage rates. Our results suggest that frequently burned grasslands are unlikely to exhibit enhanced C sequestration with increasing atmospheric CO₂ levels or N deposition.     

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].     

Elevated pCO2 Affects C and N Metabolism in Wild Type and Transgenic Tobacco Exhibiting Altered C/N Balance in Metabolite Analysis

Abstract: Diurnal changes in starch, sugar and amino acid concentrations in source leaves, sink leaves and roots of tobacco plants were determined. In addition to wild type tobacco, transformed plants deficient in root nitrate reductase and exhibiting decreased rates of growth were employed. Further, the growth rates of tobacco plants were modulated by exposure to elevated pCO₂. From the diurnal alterations in metabolite concentrations, the daily turnover of starch and amino N was estimated in order to: (i) elucidate whether turnover rates can be related to growth rates, and (ii) identify individual amino compounds with the potential to indicate nitrogen fluxes and the C/N status of plants. Elevated pCO₂ increased growth rates and daily turnover of starch in both wild type and transformed plants, indicating enhanced rates of photosynthesis. In wild type plants, elevated pCO₂ increased the turnover of amino N, notably glutamine and alanine, in mature source leaves, indicating enhanced nitrate reduction. By contrast, amino N turnover in source leaves of transformed plants was not affected by elevated pCO₂, although nitrate reduction was presumably enhanced. Apparently, export of amino N was increased from the source leaves of transformed plants. This assumption was supported by a significantly increased turnover of amino N in young sink leaves compared to mature source leaves, indicating a preference for acropetal amino N allocation and import into the young leaves of the transformed plants. Further, elevated pCO₂ increased the allocation of leaf‐derived amino N to the roots of transformed plants. This led to increased levels of amino compounds during the entire day, notably glutamate, but did not affect root growth of the transformed plants. The suitability of individual amino compounds as markers for major N fluxes, such as nitrate reduction, photorespiration, and amino N export and import is discussed.     

Net root growth and nutrient acquisition in response to predicted climate change in two contrasting heathland species

Background and aims

Accurate predictions of nutrient acquisition by plant roots and mycorrhizas are critical in modelling plant responses to climate change.

Methods

We conducted a field experiment with the aim to investigate root nutrient uptake in a future climate and studied root production by ingrowth cores, mycorrhizal colonization, and fine root N and P uptake by root assay of Deschampsia flexuosa and Calluna vulgaris.

Results

Net root growth increased under elevated CO₂, warming and drought, with additive effects among the factors. Arbuscular mycorrhizal colonization increased in response to elevated CO₂, while ericoid mycorrhizal colonization was unchanged. The uptake of N and P was not increased proportionally with root growth after 5 years of treatment.

Conclusions

While aboveground biomass was unchanged, the root growth was increased under elevated CO₂. The results suggest that plant production may be limited by N (but not P) when exposed to elevated CO₂. The species-specific response to the treatments suggests different sensitivity to global change factors, which could result in changed plant competitive interactions and belowground nutrient pool sizes in response to future climate change.     

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.     

Carbon dioxide‐ and temperature‐mediated changes in plant defensive compounds alter food utilization of herbivores

Although the impact of elevated carbon dioxide and rising temperature on plants and animals has been extensively documented recently, only limited understanding exists regarding their combined effects. The objective of this research was to address the consequences of using combinations of elevated CO₂ and elevated temperature on a plant's defensive chemistry, and subsequent utilization of the plant as insect food. Our results indicated that elevated CO₂ and increased temperature, for the most part, act independently on the production of defensive compounds in broccoli leaves (Brassica oleracea L. var. italica). CO₂ concentrations had significant effects on the foliar water content, total phenolic compounds, polyphenol oxidase and trypsin inhibitor concentrations. The herbivore Spodoptera litura (Fabricius; Lepidoptera: Noctuidae) responded to changes in the plant secondary chemistry, with larvae consuming more plant materials that had been exposed to elevated CO₂. The food utilization efficiencies of second‐instar larvae were more sensitive to CO₂‐treated foliage than those of the third‐ and fourth‐instar larvae. Temperature did exert a significant effect on food utilization (ECD) by the larvae. Our study will provide important information in future predictions on plant–insect interactions as a result of climate change. The study also demonstrated that since various larval stages might respond differently to climate change, this possibility needs to be considered in future forecasting and monitoring.     

Above- and belowground response of Populus grandidentata to elevated atmospheric CO2 and soil N availability

Soil N availability may play an important role in regulating the long-term responses of plants to rising atmospheric CO₂ partial pressure. To further examine the linkage between above- and belowground C and N cycles at elevated CO₂, we grew clonally propagated cuttings of Populus grandidentata in the field at ambient and twice ambient CO₂ in open bottom root boxes filled with organic matter poor native soil. Nitrogen was added to all root boxes at a rate equivalent to net N mineralization in local dry oak forests. Nitrogen added during August was enriched with ¹⁵N to trace the flux of N within the plant-soil system. Above-and belowground growth, CO₂ assimilation, and leaf N content were measured non-destructively over 142 d. After final destructive harvest, roots, stems, and leaves were analyzed for total N and ¹⁵N. There was no CO₂ treatment effect on leaf area, root length, or net assimilation prior to the completion of N addition. Following the N addition, leaf N content increased in both CO₂ treatments, but net assimilation showed a sustained increase only in elevated CO₂ grown plants. Root relative extension rate was greater at elevated CO₂, both before and after the N addition. Although final root biomass was greater at elevated CO₂, there was no CO₂ effect on plant N uptake or allocation. While low soil N availability severely inhibited CO₂ responses, high CO₂ grown plants were more responsive to N. This differential behavior must be considered in light of the temporal and spatial heterogeneity of soil resources, particularly N which often limits plant growth in temperate forests.     

Atmospheric CO2 enrichment and drought stress modify root exudation of barley

Rising CO₂ concentrations associated with drought stress is likely to influence not only aboveground growth, but also belowground plant processes. Little is known about root exudation being influenced by elements of climate change. Therefore, this study wanted to clarify whether barley root exudation responds to drought and CO₂ enrichment and whether this reaction differs between an old and a recently released malting barley cultivar. Barley plants were grown in pots filled with sand in controlled climate chambers at ambient (380 ppm) or elevated (550 ppm) atmospheric [CO₂] and a normal or reduced water supply. Root exudation patterns were examined at the stem elongation growth stage and when the inflorescences emerged. At both dates, root exudates were analyzed for different compounds such as total free amino acids, proline, potassium, and some phytohormones. Elevated [CO₂] decreased the concentrations in root exudates of some compounds such as total free amino acids, proline, and abscisic acid. Moreover, reduced water supply increased proline, potassium, electric conductivity, and hormone concentrations. In general, the modern cultivar showed higher concentrations of proline and abscisic acid than the old one, but the cultivars responded differentially under elevated CO₂. Plant developmental stage had also an impact on the root exudation patterns of barley. Generally, we observed significant effects of CO₂ enrichment, watering levels, and, to a lesser extent, cultivar on root exudation. However, we did not find any mitigation of the adverse effects of drought by elevated CO₂. Understanding the multitude of relationships within the rhizosphere is an important aspect that has to be taken into consideration in the context of crop performance and carbon balance under conditions of climate change.     

Effects of elevated CO2, temperature and N fertilization on nitrogen fluxes in a temperate grassland ecosystem

The soil nitrogen cycle was investigated in a pre‐established Lolium perenne sward on a loamy soil and exposed to ambient and elevated atmospheric CO₂ concentrations (350 and 700 μL L^?1) and, at elevated [CO₂], to a 3 °C temperature increase. At two levels of mineral nitrogen supply, N– (150 kgN ha^?1 y^?1) and N+ (533 kgN ha^?1 y^?1), ¹⁵N‐labelled ammonium nitrate was supplied in split applications over a 2.5‐y period. The recovery of the labelled fertilizer N was measured in the harvests, in the stubble and roots, in the macro‐organic matter fractions above 200 μm in size (MOM) and in the aggregated organic matter below 200 μM (AOM). Elevated [CO₂] reduced the total amount of N harvested in the clipped parts of the sward. The harvested N derived from soil was reduced to a greater extent than that derived from fertilizer. At both N supplies, elevated [CO₂] modified the allocation of the fertilizer N in the sward, in favour of the stubble and roots and significantly increased the recovery of fertilizer N in the soil macro‐organic matter fractions. The increase of fertilizer N immobilization in the MOM was associated with a decline of fertilizer N uptake by the grass sward, which supported the hypothesis of a negative feedback of elevated [CO₂] on the sward N yield and uptake. Similar and even more pronounced effects were observed for the native N mineralized in the soil. At N–, a greater part of the fertilizer N organized in the root phytomass resulted in an underestimation of N immobilized in dead roots and, in turn, an underestimation of N immobilization in the MOM. The 3 °C temperature increase alleviated the [CO₂] effect throughout much of the N cycle, increasing soil N mineralization, N derived from soil in the harvests, and the partitioning of the assimilated fertilizer N to shoots. In conclusion, at ambient temperature, the N cycle was slowed down under elevated [CO₂], which restricted the increase in the aboveground production of the grass sward, and apparently contributed to the sequestration of carbon belowground. In contrast, a temperature increase under elevated [CO₂] stimulated the soil nitrogen cycle, improved the N nutrition of the sward and restricted the magnitude of the soil C sequestration.     

Nitrogen to phosphorus ratios of tree species in response to elevated carbon dioxide and nitrogen addition in subtropical forests

Increased atmospheric carbon dioxide (CO₂) concentrations and nitrogen (N) deposition induced by human activities have greatly influenced the stoichiometry of N and phosphorus (P). We used model forest ecosystems in open‐top chambers to study the effects of elevated CO₂ (ca. 700 μmol mol^?1) alone and together with N addition (100 kg N ha^?1 yr^?1) on N to P (N : P) ratios in leaves, stems and roots of five tree species, including four non‐N₂ fixers and one N₂ fixer, in subtropical China from 2006 to 2009. Elevated CO₂ decreased or had no effects on N : P ratios in plant tissues of tree species. N addition, especially under elevated CO₂, lowered N : P ratios in the N₂ fixer, and this effect was significant in the stems and the roots. However, only one species of the non‐N₂ fixers showed significantly lower N : P ratios under N addition in 2009, and the others were not affected by N addition. The reductions of N : P ratios in response to elevated CO₂ and N addition were mainly associated with the increases in P concentrations. Our results imply that elevated CO₂ and N addition could facilitate tree species to mitigate P limitation by more strongly influencing P dynamics than N in the subtropical forests.