Climate change goes underground: effects of elevated atmospheric CO<Subscript>2</Subscript> on microbial community structure and activities in the rhizosphere

General concern about climate change has led to growing interest in the responses of terrestrial ecosystems to elevated concentrations of CO₂ in the atmosphere. Experimentation during the last two to three decades using a large variety of approaches has provided sufficient information to conclude that enrichment of atmospheric CO₂ may have severe impact on terrestrial ecosystems. This impact is mainly due to the changes in the organic C dynamics as a result of the effects of elevated CO₂ on the primary source of organic C in soil, i.e., plant photosynthesis. As the majority of life in soil is heterotrophic and dependent on the input of plant-derived organic C, the activity and functioning of soil organisms will greatly be influenced by changes in the atmospheric CO₂ concentration. In this review, we examine the current state of the art with respect to effects of elevated atmospheric CO₂ on soil microbial communities, with a focus on microbial community structure. On the basis of the existing information, we conclude that the main effects of elevated atmospheric CO₂ on soil microbiota occur via plant metabolism and root secretion, especially in C3 plants, thereby directly affecting the mycorrhizal, bacterial, and fungal communities in the close vicinity of the root. There is little or no direct effect on the microbial community of the bulk soil. In particular, we have explored the impact of these changes on rhizosphere interactions and ecosystem processes, including food web interactions.     

Effect of elevated atmospheric CO2 concentration on growth and physiology of wheat and sorghum under cadmium stress

ABSTRACT

Elevated concentrations of carbon dioxide (e[CO₂]) affect plant growth and physiological characteristics, including metal accumulation, and the activity of anti-oxidant enzymes. These effects were investigated in cadmium (Cd) tolerant wheat (Triticum aestivum L.) and sorghum (Sorghum bicolor (L.) Moench.) cultivars. Plants were grown at the ambient and elevated CO₂ levels, with four concentrations of Cd (0, 10, 20 and 40 mg kg^?1) added to the soil. After 60 days, subsamples were tested for chlorophylls and carotenoids, protein, enzyme activities and morphological characteristics.

Results showed that e[CO₂] increased plant height, leaf area, and the dry weight of shoots and roots (P < 0.01). In addition, it decreased the Cd concentration in the shoots and roots of wheat, and increased the same concentrations for sorghum. With increasing Cd, the activities of the anti-oxidants, SOD and GSH-px increased in wheat. The differences in enzyme activity parallel the changes in Cd concentration in the plants of both species.     

Effect of elevated carbon dioxide and water stress on gas exchange and water use efficiency in corn

CO₂ has been predicted to increase in the future, and thus leading to possible changes in precipitation patterns. The objectives of this study were to investigate water use and canopy level photosynthesis of corn plants, and to quantify water use efficiency in corn plants under two different CO₂ levels combined with four different water stress levels. Corn plants were planted in sunlit plant growth chambers and a day/night temperature of (28/18 °C) was applied. From 21 days after emergence (DAE), the eight treatments including two levels of carbon dioxide concentrations (400 and 800 μmol mol⁻¹) and four levels of water stress (well-watered control, “mild”, “moderate”, and “severe” water stress) treatments at each CO₂ level were imposed. Height, number of leaves, leaf lengths, and growth stages of corn plants were monitored from nine plants twice a week. Corn plants were separately collected, dried, and analyzed for the biomass accumulation at 21 and 60 DAE. Soil water contents were monitored by a time domain reflectometry (TDR) system (15 probes per chamber). The “breaking points” (changes from high to low rates of soil water uptake) were observed in the bottom of soil depth for the water stressed conditions, and the “breaking points” under ambient CO₂ appeared 6-9 days earlier than under elevated CO₂. Although approximately 20-49% less water was applied for the elevated CO₂ treatments than for ambient CO₂ from 21 DAE, higher soil water contents were recorded under elevated CO₂ than under ambient CO₂. However, corn growth variables such as height, leaf area, and biomass accumulation were not significantly different in CO₂ or water stressed treatments. This result may be explained by considering that significant differences in canopy level gross photosynthesis among the water stress treatments was observed only toward the end of the experiment. The higher soil water contents observed under elevated CO₂ resulted mainly from less water use than under ambient CO_2. WUE (above ground biomass per water use since 21 DAE) at the final harvest was consistently higher and varied with a smaller range under elevated CO₂ than under ambient CO₂. This study suggests that less water will be required for corn under high-CO₂ environment in the future than at present.     

CO2浓度升高对不同水分条件下冬小麦生长和水分利用的影响

以CO₂浓度升高为主要特征的气候变化对作物生长发育及产量形成的影响日益受到重视。冬小麦是我国主要粮食作物之一, 主要分布在干旱及半干旱地区, 且生长期内多干旱少雨。研究不同水分条件下冬小麦的生长变化及水分利用对CO₂浓度升高的响应具有重要的科学和实践意义。本研究在封顶式生长室中对2个土壤水分水平[适宜水分: 70%~80%田间持水量; 干旱胁迫: 50%~60%田间持水量]的盆栽冬小麦进行了CO₂熏蒸试验[背景大气浓度: (396.1±29.2) μmol·mol^-1; 升高的浓度: (760.1±36.1)μmol·mol^-1]。对小麦植株生理指标、生物量、产量、耗水量和水分利用效率(WUE)等的研究结果表明, 与背景大气CO₂浓度相比, CO₂浓度升高可促进冬小麦生长, 其地上生物量显著增加, 适宜水分和干旱胁迫条件下分别增加了28.6%和18.6%; 籽粒产量显著增加, 适宜水分和干旱胁迫条件下分别增加了32.6%和22.6%; CO₂浓度升高主要通过增加穗粒数提高籽粒产量, 穗粒数在适宜水分条件下提高24.3%, 干旱胁迫条件下提高15.5%, 对千粒重没有显著影响。CO₂浓度升高使群体和产量WUE显著提高, 在适宜水分条件下提高幅度较大, 分别提高17.7%和24.8%。CO₂浓度升高显著提高了叶片光合速率(P_n)、降低了气孔导度(G_s)和蒸腾速率(T_r); 在适宜水分和干旱胁迫下P_n分别提高15.6%与12.9%, G_s分别降低22.7%与18.2%, T_r分别降低8.9%与7.5%。CO₂浓度升高提高了叶片水势及叶绿素含量; 在适宜水分条件下叶片水势提高幅度较大, 为7.7%; 叶片叶绿素含量在2种水分条件分别提高7.5%与3.8%。由以上试验结果可得出: CO₂浓度升高对冬小麦的生长、产量及水分利用效率均具有促进作用, 而且在土壤水分状况较好时, 这种作用效果更明显; CO₂浓度升高主要通过增加穗粒数来促进产量提高。     

Impact of elevated CO2 on mycorrhizal associations and implications for plant growth

The impact of increasing concentrations of atmospheric CO₂ upon plant physiology has been widely investigated. Plant, and in particular root, growth is nearly always enhanced as a direct consequence of CO₂ enrichment, with C₃ species generally more responsive than C₄ species. Such alterations in plant productivity will have consequence for below-ground processes and increased carbon allocation to the roots may favour symbiotic relationships. This paper discusses the current information available for the consequences of these changes upon mycorrhizal relationships. Generally mycorrhizal plants grown under CO₂ enrichment show enhanced phosphorus uptake but nitrogen uptake is unaffected. This increased nutrient uptake is not correlated with increased mycorrhizal colonization of the roots. Similarly root exudation does not increase under CO₂ enrichment but qualitative differences have yet to be assessed. However, it is predicted that total rhizodeposition of materials will increase as will litter inputs, although mineral and biochemical alterations to these plant derived inputs may occur. The consequences of such changes within the rhizosphere are discussed and future research     

Elevated CO2 increases the effect of an arbuscular mycorrhizal fungus and a plant-growth-promoting rhizobacterium on structural stability of a semiarid agricultural soil under drought conditions

In arid and semiarid Mediterranean regions, an increase in the severity of drought events could be caused by rising atmospheric CO₂ concentrations. We studied the effects of the interaction of CO₂, water supply and inoculation with a plant-growth-promoting rhizobacterium (PGPR), Pseudomonas mendocina Palleroni, or inoculation with an arbuscular mycorrhizal (AM) fungus, Glomus intraradices (Schenk & Smith), on aggregate stabilisation of the rhizosphere soil of Lactuca sativa L. cv. Tafalla. The influence of such structural improvements on the growth of lettuce was evaluated. We hypothesised that elevated atmospheric CO₂ concentration would increase the beneficial effects of inoculation with a PGPR or an AM fungus on the aggregate stability of the rhizosphere soil of lettuce plants. Leaf hydration, shoot dry biomass and mycorrhizal colonisation were decreased significantly under water-stress conditions, but this decrease was more pronounced under ambient vs elevated CO₂. The root biomass decreased under elevated CO₂ but only in non-stressed plants. Under elevated CO₂, the microbial biomass C of the rhizosphere of the G. intraradices-colonised plants increased with water stress. Bacterial and mycorrhizal inoculation and CO₂ had no significant effect on the easily-extractable glomalin concentration. Plants grown under elevated CO₂ had a significantly higher percentage of stable aggregates under drought stress than under well-watered conditions, particularly the plants inoculated with either of the assayed microbial inocula (about 20% higher than the control soil). We conclude that the contribution of mycorrhizal fungi and PGPR to soil aggregate stability under elevated atmospheric CO₂ is largely enhanced by soil drying.     

Rhizodeposition-induced decomposition increases N availability to wild and cultivated wheat genotypes under elevated CO2

Elevated CO₂ may increase nutrient availability in the rhizosphere by stimulating N release from recalcitrant soil organic matter (SOM) pools through enhanced rhizodeposition. We aimed to elucidate how CO₂-induced increases in rhizodeposition affect N release from recalcitrant SOM, and how wild versus cultivated genotypes of wheat mediated differential responses in soil N cycling under elevated CO₂. To quantify root-derived soil carbon (C) input and release of N from stable SOM pools, plants were grown for 1 month in microcosms, exposed to ¹³C labeling at ambient (392 μmol mol⁻¹) and elevated (792 μmol mol⁻¹) CO₂ concentrations, in soil containing ¹⁵N predominantly incorporated into recalcitrant SOM pools. Decomposition of stable soil C increased by 43%, root-derived soil C increased by 59%, and microbial-¹³C was enhanced by 50% under elevated compared to ambient CO₂. Concurrently, plant ¹⁵N uptake increased (+7%) under elevated CO₂ while ¹⁵N contents in the microbial biomass and mineral N pool decreased. Wild genotypes allocated more C to their roots, while cultivated genotypes allocated more C to their shoots under ambient and elevated CO₂. This led to increased stable C decomposition, but not to increased N acquisition for the wild genotypes. Data suggest that increased rhizodeposition under elevated CO₂ can stimulate mineralization of N from recalcitrant SOM pools and that contrasting C allocation patterns cannot fully explain plant mediated differential responses in soil N cycling to elevated CO₂.     

Effects of elevated CO2 concentration on rhizodeposition from Lolium perenne grown on soil exposed to 9 years of CO2 enrichment

The effects of enriched CO₂ atmosphere on partitioning of recently assimilated carbon were investigated in a plant-soil-microorganism system in which Lolium perenne seedlings were planted into cores inserted into the resident soil within a sward that had been treated with elevated CO₂ for 9 consecutive years, under two N fertilisation levels (Swiss FACE experiment). The planted cores were excavated from the ambient (35 Pa pCO₂) and enriched (60 Pa pCO₂) rings at two dates, in spring and autumn, during the growing season. The cores were brought back to the laboratory for ¹⁴C labelling of shoots in order to trace the transfer of recently assimilated C both within the plant and to the soil and microbial biomass. At the spring sampling, high N supply stimulated shoot and total dry matter production. Consistently, high N enhanced the allocation of recently fixed C to shoots, and reduced it to belowground compartments. Elevated CO₂ had no consequences for DM or the pattern of C allocation. At the autumn sampling, at high N plot, yield of L. perenne was stimulated by elevated CO₂. Consistently, ¹⁴C was preferentially allocated aboveground and, consequently belowground recent C allocation was depressed and rhizodeposition reduced. At both experimental periods, total soil C content was similar in all treatments, providing no evidence for soil carbon sequestration in the Swiss Free Air CO₂ Enrichment experiment (FACE) after 9 years of enrichment. Recently assimilated C and soil C were mineralised faster in soils from enriched rings, suggesting a CO₂-induced shift in the microbial biomass characteristics (structure, diversity, activity) and/or in the quality of the root-released organic compounds.     

Increased moisture and methanogenesis contribute to reduced methane oxidation in elevated CO2 soils

Awareness of global warming has stimulated research on environmental controls of soil methane (CH₄) consumption and the effects of increasing atmospheric carbon dioxide (CO₂) on the terrestrial CH₄ sink. In this study, factors impacting soil CH₄ consumption were investigated using laboratory incubations of soils collected at the Free Air Carbon Transfer and Storage I site in the Duke Forest, NC, where plots have been exposed to ambient (370 μL L⁻¹) or elevated (ambient + 200 μL L⁻¹) CO₂ since August 1996. Over 1 year, nearly 90% of the 360 incubations showed net CH₄ consumption, confirming that CH₄-oxidizing (methanotrophic) bacteria were active. Soil moisture was significantly (p < 0.01) higher in the 25–30 cm layer of elevated CO₂ soils over the length of the study, but soil moisture was equal between CO₂ treatments in shallower soils. The increased soil moisture corresponded to decreased net CH₄ oxidation, as elevated CO₂ soils also oxidized 70% less CH₄ at the 25–30 cm depth compared to ambient CO₂ soils, while CH₄ consumption was equal between treatments in shallower soils. Soil moisture content predicted (p < 0.05) CH₄ consumption in upper layers of ambient CO₂ soils, but this relationship was not significant in elevated CO₂ soils at any depth, suggesting that environmental factors in addition to moisture were influencing net CH₄ oxidation under elevated CO₂. More than 6% of the activity assays showed net CH₄ production, and of these, 80% contained soils from elevated CO₂ plots. In addition, more than 50% of the CH₄-producing flasks from elevated CO₂ sites contained deeper (25–30 cm) soils. These results indicate that subsurface (25 cm+) CH₄ production contributes to decreased net CH₄ consumption under elevated CO₂ in otherwise aerobic soils.     

Seed priming to mitigate the impact of elevated carbon dioxide associated temperature stress on germination in rice (Oryza sativa L.)

ABSTRACT

Climate change has a negative impact on crop production by inducing several stresses throughout plant growth. As the germination is one of the critical stages, this study was to assess the impact of elevated carbon dioxide (eCO₂) and associated temperature stress on germination of rice. The study was conducted using the rice genotype CO 51 in artificially induced eCO₂ condition using open-top chamber (OTC). The efficiency of seed priming treatment with salicylic acid (SA), citric acid (CA), ascorbic acid (AsA) and distilled water (hydropriming) to alleviate stress due to eCO₂ and temperature were also studied. SA 25 mg l^?1 and AsA 100 mg l^?1 enhanced germination, other seed quality parameters along with α- amylase activity. Also, the activity of antioxidant enzymes like catalase, peroxidase and superoxide dismutase was increased. These parameters positively affected the germination and growth by mitigating the effect of oxidative stress induced under eCO₂ conditions. So, SA, CA and AsA can be effectively used for maintaining seed quality parameters and seedling growth during the stresses.     

Evidence of N<Subscript>2</Subscript>O emission and gaseous nitrogen losses through nitrification-denitrification induced by rice plants (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

A greenhouse experiment was conducted with a specially designed apparatus consisting of an upper and lower chamber where the treatment with rice was carried out (treatment 1). The apparatus also had a single chamber where treatment 2, without rice plants, was carried out. The scope of this study was to elucidate the influence of rice plant growth on gaseous N losses as N₂ and N₂O produced by nitrification-denitrification in a flooded soil fertilized with (NH₄)₂SO₄ (with 56.50 atom% ¹⁵N). Gas samples were withdrawn weekly and analyzed for (N₂ + N₂O)-¹⁵N losses by mass spectrometer and for N₂O by gas chromatograph. The gaseous (N₂ + N₂O)-¹⁵N losses of the treatment with rice plants were significantly (P =0.01) higher than those of the treatment without rice plants, as were the amounts of N₂O emitted. Rice plants facilitate the efflux of N₂ and N₂O from soil to atmosphere, as about half of the total gaseous ¹⁵N loss as N₂ and N₂O was found in the upper chamber. The proportion of N₂O-¹⁵N to (N₂ + N₂O)-¹⁵N in the upper chamber was 10.56%, much higher than that of the lower chamber in treatment 1 and the headspace of treatment 2.     

The counteracting effects of elevated atmospheric CO2 concentrations and drought episodes: Studies of enchytraeid communities in a dry heathland

The potential impacts of interactions of multiple climate change factors in soil ecosystems have received little attention. Most studies have addressed effects of single factors such as increased temperature or atmospheric CO₂ but little is known about how such environmental factors will interact. In the present study we investigate the effects of in situ exposure to elevated atmospheric CO₂ concentration, increased temperatures and prolonged drought episodes on field communities of Enchytraeidae (Oligochaeta) in a dry heathland (Brandbjerg, Denmark). Increased CO₂ had a positive effect on enchytraeid biomass, whereas drought significantly reduced it. Elevated temperature did not result in any detectable effects. No interactions between the three factors were observed. Interestingly, the positive effect of increased CO₂ and the negative effect of drought were cancelled out when applied in combination. Thus, in the combined drought and CO₂ treatment, and when additionally combined with increased temperature, the total biomass of enchytraeids was similar to those in the ambient plots. The positive effect of increased CO₂ seemed to be influenced by season, as the response was most pronounced during summer, and less so in autumn. The impact of the drought was more consistent, with enchytraeids reduced at all sampling occasions. Moreover, the negative effect of drought seemed to depend on the inter-annual variability of precipitation. The year with a dry summer and autumn (2006) showed a stronger impact of drought on the enchytraeids, compared to the year with a wet summer and autumn (2007). Our study emphasises the importance of multi-factorial experimental design as a means to investigate effects of climatic changes.     

Denitrification in grass swards is increased under elevated atmospheric CO2

Emissions of N₂O and N₂ were measured from Lolium perenne L. swards under ambient (36 Pa) and elevated (60 Pa) atmospheric CO₂ at the Swiss free air carbon dioxide enrichment experiment following application of 11.2 g N m⁻² as ¹⁵NH₄¹⁵NO₃ or ¹⁴NH₄¹⁵NO₃ (1 at.% excess ¹⁵N). Total denitrification (N₂O+N₂) was increased under elevated pCO₂ with emissions of 6.2 and 19.5 mg ¹⁵N m⁻² measured over 22 d from ambient and elevated pCO₂ swards, respectively, supporting the hypothesis that increased belowground C allocation under elevated pCO₂ provides the energy for denitrification. Nitrification was the predominant N₂O producing process under ambient pCO₂ whereas denitrification was predominant under elevated pCO₂. The N₂-to-N₂O ratio was often higher under elevated pCO₂ suggesting that previous estimates of gaseous N losses based only on N₂O emissions have greatly underestimated the loss of N by denitrification.     

大气CO2浓度升高和干旱互作对谷子光合及抗旱生理特性的影响

为明确谷子光合作用以及抗旱生理过程对高大气CO_2浓度和干旱交互作用的响应机制,在开顶式气室中(OTC)开展大气CO_2浓度和干旱交互对谷子影响的研究。设置两个CO_2浓度:环境CO_2浓度(400μmol·mol~(-1))和高CO_2浓度(600μmol·mol~(-1));两个水分处理:正常水分(70%～80%田间持水量)和干旱(45%～55%田间持水量),对高CO_2浓度和干旱互作下谷子光合气体交换参数、荧光动力学参数及抗旱相关生理指标的变化进行了研究。结果表明:高CO_2浓度可降低干旱条件下光合色素含量,加剧孕穗期谷子气孔关闭,减轻灌浆期干旱对谷子净光合速率的负效应并增加其水分利用效率。孕穗期高CO_2处理使正常水分处理下谷子气孔导度下降66.7%,而干旱处理下减少77.7%;灌浆期高CO_2使正常水分处理和干旱处理下谷子净光合速率分别增加19.0%和87.7%,水分利用效率增加37.1%和39.2%。干旱处理显著降低谷子除非光化学淬灭系数(NPQ)以外所有荧光动力学参数值,灌浆期高CO_2能缓解该作用。高CO_2处理显著减少纤维素含量和正常水分处理下过氧化物酶活性。干旱极显著升高POD活性(高CO_2浓度)及脯氨酸含量、可溶性总糖、淀粉含量(环境CO_2浓度)和纤维素含量(高CO_2浓度)。因此CO_2浓度升高能够改善谷子的PSⅡ光化学效率和提高抗氧化酶活性来增强谷子的抗旱性。     

Palatability trials on hardwood leaf litter grown under elevated CO2: a stable carbon isotope study

The palatability to isopods and microbes of a broad range of hardwood leaf litter, derived from three field CO₂-enrichment experiments in the USA, was investigated, using δ¹³C, to trace the C flow from litter to isopods and to CO₂ respired by microbial decomposition. Leaf litter grown under elevated CO₂ had δ¹³C values ranging from −39 to −45‰, which were significantly different from ambient litter δ¹³C values of around −30‰. Litter palatability to isopods of the Porcellio sp. was tested by incubating ambient- and elevated-CO₂ litter, and a mixture of the two, in the presence of isopods for 14 days, under environmentally controlled conditions; δ¹³C was measured on litter and isopods' body before and after incubation. In an additional experiment, litter was incubated in the absence of fauna for 30 days, and on five occasions the δ¹³C of the CO₂ respired from litter was measured. The ¹³C label was clearly carried from the litter source to the isopods' bodies, and their faeces. For microbial-respired CO₂, δ¹³C was significantly higher than that of the litter source, suggesting preferential degradation of substrates enriched in ¹³C as compared to those in the overall litter. With the exception of Quercus myrtifolia leaf litter, elevated CO₂ did not affect the palatability to isopods nor the microbial degradation of any of the litters, possibly as a result of unaltered litter N concentration. However, significant differences in litter palatability and decay rates were observed among the different species. With this study, the use of isotopically labelled litter material was confirmed as a key methodology that can significantly contribute to the advancement of the understanding of litter decomposition and of the quantification of C fluxes in the process.     

Nitrogen fixation by biological soil crusts and heterotrophic bacteria in an intact Mojave Desert ecosystem with elevated CO2 and added soil carbon

Fixation of N by biological soil crusts and free-living heterotrophic soil microbes provides a significant proportion of ecosystem N in arid lands. To gain a better understanding of how elevated CO₂ may affect N₂-fixation in aridland ecosystems, we measured C₂H₂ reduction as a proxy for nitrogenase activity in biological soil crusts for 2 yr, and in soils either with or without dextrose-C additions for 1 yr, in an intact Mojave Desert ecosystem exposed to elevated CO₂. We also measured crust and soil δ¹⁵N and total N to assess changes in N sources, and δ¹³C of crusts to determine a functional shift in crust species, with elevated CO₂. The mean rate of C₂H₂ reduction by biological soil crusts was 76.9±5.6 μmol C₂H₄ m⁻² h⁻¹. There was no significant CO₂ effect, but crusts from plant interspaces showed high variability in nitrogenase activity with elevated CO₂. Additions of dextrose-C had a positive effect on rates of C₂H₂ reduction in soil. There was no elevated CO₂ effect on soil nitrogenase activity. Plant cover affected soil response to C addition, with the largest response in plant interspaces. The mean rate of C₂H₂ reduction in soils either with or without C additions were 8.5±3.6 μmol C₂H₄ m⁻² h⁻¹ and 4.8±2.1 μmol m⁻² h⁻¹, respectively. Crust and soil δ¹⁵N and δ¹³C values were not affected by CO₂ treatment, but did show an effect of cover type. Crust and soil samples in plant interspaces had the lowest values for both measurements. Analysis of soil and crust [N] and δ¹⁵N data with the Rayleigh distillation model suggests that any plant community changes with elevated CO₂ and concomitant changes in litter composition likely will overwhelm any physiological changes in N₂-fixation.     

Effects of elevated CO2 and O3 on soil respiration under ponderosa pine

Soil respiration represents the integrated response of plant roots and soil organisms to environmental conditions and the availability of C in the soil. A multi-year study was conducted in outdoor sun-lit controlled-environment chambers containing a reconstructed ponderosa pine/soil-litter system. The study used a 2×2 factorial design with two levels of CO₂ and two levels of O₃ and three replicates of each treatment. The objectives of our study were to assess the effects of long-term exposure to elevated CO₂ and O₃, singly and in combination, on soil respiration, fine root growth and soil organisms. Fine root growth and soil organisms were included in the study as indicators of the autotrophic and heterotrophic components of soil respiration. The study evaluated three hypotheses: (1) elevated CO₂ will increase C assimilation and allocation belowground increasing soil respiration; (2) elevated O₃ will decrease C assimilation and allocation belowground decreasing soil respiration and (3) as elevated CO₂ and O₃ have opposing effects on C assimilation and allocation, elevated CO₂ will eliminate or reduce the negative effects of elevated O₃ on soil respiration. A mixed-model covariance analysis was used to remove the influences of soil temperature, soil moisture and days from planting when testing for the effects of CO₂ and O₃ on soil respiration. The covariance analysis showed that elevated CO₂ significantly reduced the soil respiration while elevated O₃ had no significant effect. Despite the lack of a direct CO₂ stimulation of soil respiration, there were significant interactions between CO₂ and soil temperature, soil moisture and days from planting indicating that elevated CO₂ altered soil respiration indirectly. In elevated CO₂, soil respiration was more sensitive to soil temperature changes and less sensitive to soil moisture changes than in ambient CO₂. Soil respiration increased more with days from planting in elevated than in ambient CO₂. Elevated CO₂ had no effect on fine root biomass but increased abundance of culturable bacteria and fungi suggesting that these increases were associated with increased C allocation belowground. Elevated CO₂ had no significant effect on microarthropod and nematode abundance. Elevated O₃ had no significant effects on any parameter except it reduced the sensitivity of soil respiration to changes in temperature.     

C and N in soil organic matter density fractions under elevated atmospheric CO2: Turnover vs. stabilization

Turnover of C and N in an arable soil under Free Air Carbon Dioxide (FACE) experiment was studied by the use of ¹³C natural abundance and ¹⁵N-labeled fertilizers. Wheat was kept four growing seasons under ambient and elevated CO₂ concentrations and fertilized for three growing seasons. Density fractionation of soil organic matter (SOM) allowed to track ¹³C and ¹⁵N in free particulate organic matter (fPOM; <1.6 g cm⁻³), particulate organic matter occluded within aggregates with two densities (oPOM 1.6, oPOM 1.6-2.0 g cm⁻³), and in mineral-associated organic matter (>2.0 g cm⁻³) fractions. Elevated CO₂ and N fertilization did not significantly affect C and N contents in the bulk soil. Calculated mean residence time (MRT) of C and N revealed the qualitative differences of SOM density fractions: (i) the shortest MRT_C and MRT_N in fPOM confirmed high availability of this fraction to decomposition. Larger C/N ratio of fPOM under elevated vs. ambient CO₂ indicated an increasing recalcitrance of FACE-derived plant residues. (ii) There was no difference in MRT of C and N between lighter and heavier oPOMs probably due to short turnover time of soil aggregates which led to oPOM mixing. The increase of MRT_C and MRT_N in both oPOMs during the experiment confirmed the progressive degradation of organic material within aggregates. (iii) Constant turnover rates of C in the mineral fraction neither confirmed nor rejected the assumed stabilization of SOM to take place in the mineral fraction. Moreover, a trend of decreasing of C and N amounts in the Min fraction throughout the experiment was especially pronounced for C under elevated CO₂. Hence, along with the progressive increase of C_FACE in the Min fraction the overall losses of C under elevated CO₂ may occur at the expense of older “pre-FACE” C.     

Developmental patterns of lamellae and stroma fractions of chloroplasts in rice plants

On examining the changes in lamellae and stroma nitrogen during leaf development, it is demonstrated that the lamellae and stroma fractions ofrice chloroplasts develop in quite different ways. In the case of stroma, the stroma materials existing in the leaf section which has just emerged from a leaf sheath are quite limited and the major part of this fraction is derived from the successive protein synthesis, i.e., the synthesis of this fraction was markedly increased during leaf expansion. This developmental pattern of the stroma coincided with the changes in the high-molecular-weight water soluble leaf protein, which seemed to be mainly composed of Fraction I protein. A rapid increase in stroma nitrogen was found to be a major cause for an increase in the leaf nitrogen content during leaf development.

On the other hand, the developmental pattern of the lamellae fraction was characterized by the fact that a considerable amount of this fraction had already been prepared when a leaf emerged from a leaf sheath and thereafter, no outstanding increase was seen compared to that of the stroma. This developmental pattern of the lamellae fraction resulted in a lowering of the proportion of lamellae nitrogen to the total leaf nitrogen during leaf development.

A great change in the lamellae-stroma composition of chloroplasts was observed. The proportion of stroma nitrogen to the total chloroplast nitrogen tended to increase as a leaf develops. Since the developmental stage varied according to the regions of a leaf, variation of the lamellaestroma composition was seen even within a leaf, i.e., the proportion of stroma nitrogen increased from base to tip.

In order to compare the synthetic rate of chlorophyll with those of the stroma and lamellae fractions, the changes in the ratios of stroma nitrogen/chlorophyll and lamellae nitrogen/chlorophyll were examined. The lamellae nitrogen/chlorophyll ratio decreased as a leaf developed, whereas the stroma nitrogen/chlorophyll ratio increased. Then the synthetic rates of these fractions during leaf development turned out to be of the same order as the stroma fraction, chlorophyll, lamellae fraction.     

Nitrogen Transformation as Influenced by Soil Microbial Nitrogen under Carbon Dioxide Enrichment Induced in Created Riparian Wetlands

Limited research has been conducted on how atmospheric carbon dioxide (CO₂) affects water and soil nitrogen (N) transformation in wetland ecosystems. A stable isotope technique is suitable for conducting a detailed investigation of mechanistic nutrient transformations. Nutrient ammonium sulfate (NH₄)₂SO₄ input in culture water under elevated CO₂ (700 μL L^?1) and ambient CO₂ (380 μL L^?1) was studied to analyze N transformations with N blanks for both water and soil. It was measured by ¹⁵N pool dilution using analytical equations in a riparian wetland during a 3-month period. Soil gross ammonium (NH₄ ⁺) mineralization and consumption rates increased significantly by 22% and 404%, Whereas those of water decreased greatly by??57% and??57% respectively in enriched CO₂. In contrast, gross nitrate (NO₃ ^?) consumption and nitrification rates of soil decreased by??11% and??14% and those of water increased by 29% and 27% respectively in enrichment CO₂. These may be due to the extremely high soil microbial biomass nitrogen (MBN), which increased by 94% in elevated soil. The results can show when CO₂ concentrations are going to rise in the future. Consequently soil microbial activity initiates the decreased N concentration in sediment and increased N concentration in overlying water in riparian wetland ecosystems.