The effect of plant material grown under elevated CO2 on soil respiratory activity

The biodegradability of aerial material from a C4 plant, sorghum grown under ambient (345 µmol mol^–1) and elevated (700 µmol mol^–1) atmospheric CO₂ concentrations were compared by measuring soil respiratory activity. Initial daily respiratory activity (measured over 10 h per day) increased four fold from 110 to 440 cm³ CO₂ 100g dry weight soil^–1 in soils amended with sorghum grown under either elevated or ambient CO₂. Although soil respiratory activity decreased over the following 30 days, respiration remained significantly higher (t-test;p>0.05) in soils amended with sorghum grown under elevated CO₂ concentrations. Analysis of the plant material revealed no significant differences in C:N ratios between sorghum grown under elevated or ambient CO₂. The reason for the differences in soil respiratory activity have yet to be elucidated. However if this trend is repeated in natural ecosystems, this may have important implications for C and N cycling.     

Soil microbiota in two annual grasslands: responses to elevated atmospheric CO2

We measured soil bacteria, fungi, protozoa, nematodes, and biological activity in serpentine and sandstone annual grasslands after 4 years of exposure to elevated atmospheric CO₂. Measurements were made during the early part of the season, when plants were in vegetative growth, and later in the season, when plants were approaching their maximum biomass. In general, under ambient CO₂, bacterial biomass, total protozoan numbers, and numbers of bactivorous nematodes were similar in the two grasslands. Active and total fungal biomasses were higher on the more productive sandstone grassland compared to the serpentine. However, serpentine soils contained nearly twice the number of fungivorous nematodes compared to the sandstone, perhaps explaining the lower standing crop of fungal biomass in the serpentine and suggesting higher rates of energy flow through the fungal-based soil food web. Furthermore, root biomass in the surface soils of these grasslands is comparable, but the serpentine contains 6 times more phytophagous nematodes compared to the sandstone, indicating greater below-ground grazing pressure on plants in stressful serpentine soils. Elevated CO₂ increased the biomass of active fungi and the numbers of flagellates in both grasslands during the early part of the season and increased the number of phytophagous nematodes in the serpentine. Elevated CO₂ had no effect on the total numbers of bactivorous or fungivorous nematodes, but decreased the diversity of the nematode assemblage in the serpentine at both sampling dates. Excepting this reduction in nematode diversity, the effects of elevated CO₂ disappeared later in the season as plants approached their maximum biomass. Elevated CO₂ had no effect on total and active bacterial biomass, total fungal biomass, or the total numbers of amoebae and ciliates in either grassland during either sampling period. However, soil metabolic activity was higher in the sandstone grassland in the early season under elevated CO₂, and elevated CO₂ altered the patterns of use of individual carbon substrates in both grasslands at this time. Rates of substrate use were also significantly higher in the sandstone, indicating increased bacterial metabolic activity. These changes in soil microbiota are likely due to an increase in the flux of carbon from roots to soil in elevated CO₂, as has been previously reported for these grasslands. Results presented here suggest that some of the carbon distributed below ground in response to elevated CO₂ affects the soil microbial food web, but that these effects may be more pronounced during the early part of the growing season.     

Ecosystem water fluxes for two grasslands in elevated CO2: a modeling analysis

The need to combine data from CO₂ field experiments with climate data remains urgent, particularly because each CO₂ experiment cannot run for decades to centuries. Furthermore, predictions for a given biome need to take into account differences in productivity and leaf area index (LAI) independent of CO₂-derived changes. In this study, we use long-term weather records and field data from the Jasper Ridge CO₂ experiment in Palo Alto, California, to model the effects of CO₂ and climate variability on ecosystem water fluxes. The sandstone and serpentine grasslands at Jasper Ridge provide a range of primary productivity and LAI, with the sandstone as the more productive system. Modeled soil water availability agreed well with published observations of time-domain reflectometry in the CO₂ experiment. Simulated water fluxes based on 10-year weather data (January 1985–December 1994) showed that the sandstone grassland had a much greater proportion of water movement through plants than did the serpentine; transpiration accounted for approximately 30% of annual fluxes in the sandstone and only 10% in the serpentine. Although simulated physiological and biomass changes were similar in both grasslands, the consequences of elevated CO₂ were greater for the sandstone water budget. Elevated CO₂ increased soil drainage by 20% in the sandstone, despite an approximately one-fifth increase in plant biomass; in the serpentine, drainage increased by <10% and soil evaporation was unchanged for the same simulated biomass change. Phenological changes, simulated by a 15-day lengthening of the growing season, had minimal impacts on the water budget. Annual variation in the timing and amount of rainfall was important for water fluxes in both grasslands. Elevated CO₂ increased sandstone drainage >50 mm in seven of ten years, but the relative increase in drainage varied from 10% to 300% depending on the year. Early-season transpiration in the sandstone decreased between 26% and 41%, with elevated CO₂ resulting in a simulated water savings of 54–76 mm. Even in years when precipitation was similar (e.g., 505 and 479 mm in years 3 and 4), the effect of CO₂ varied dramatically. The response of grassland water budgets to CO₂ depends on the productivity and structure of the grassland, the amount and timing of rainfall, and CO₂-induced changes in physiology. In systems with low LAI, large physiological changes may not necessarily alter total ecosystem water budgets dramatically. Received: 11 March 1997 / Accepted: 23 September 1997     

A meta-analytical test of elevated CO2 effects on plant respiration

Contrasting results regarding elevated CO₂ effects on leafdark respiration (Rd) have hampered efforts to incorporate this importantcomponent of the plant carbon budget into long-term predictions of ecologicalresponses to rising atmospheric CO₂. To help resolve some of theseinconsistencies in the literature, we used meta-analysis to quantitativelysummarize 45 area-based leaf Rd (Rd_a) and 44 mass-based leaf Rd(Rd_m) observations from independent studies on 33 species. Ouranalysis showed that across all studies, leaf Rd_m was significantlyreduced (–18%, P < 0.05), while leaf Rd_a was marginallyincreased (+8%, P < 0.15), under elevated CO₂. There weresignificant differences among categorical groups in CO₂ effects onleaf Rd_a and Rd_m. For example, leaf Rd_a ofherbaceous species increased 28%, but leaf Rd_a of woody speciesremained unchanged under elevated CO₂. Plants exposed to elevatedCO₂ for < 60 days had significantly higher leaf Rd_a atelevated compared to ambient CO₂, while plants exposed to elevatedCO₂ for longer period of time showed no response. The magnitude ofreduction in leaf Rd_m for plants exposed to elevated CO₂for > 100 days was significantly greater than that for plants exposed toelevated CO₂ for < 100 days. Our meta-analysis of publishedresults suggest that the amount of carbon loss through leaf Rd will likelyincrease in a higher CO₂ environment because of higher leafRd_a and a proportionally greater leaf biomass increase than leafRd_m reduction at elevated CO₂. Our results alsodemonstrated the strong dependency of Rd responses to elevated CO₂onexperimental conditions.     

大气氮沉降影响草地植物物种多样性机制研究综述

大气氮沉降对草地生态系统结构和功能的影响已成为全球变化生物学研究重点。大气氮沉降导致草地群落物种多样性降低已成为全球普遍现象,但其生物学机制还不清楚,因此有必要系统梳理大气氮沉降对全球不同草地生态系统的研究结果,以便在氮沉降背景下为我国草地生态系统的研究和管理制定科学决策。系统综述了氮沉降降低草地群落物种多样性的可能机制,主要包括资源竞争排斥、群落更新限制、土壤酸化及其离子毒害、养分失衡、氮素本身的毒害、次生胁迫。氮沉降导致草地物种多样性降低是多种机制综合作用的结果,每种机制在不同时空具有不同的相对贡献。同时,与欧洲酸性土壤草地和美国高草草原相比,我国草地土壤类型和植被属性具有明显差异。因此,应根据我国草地生态系统的特征、不同植物功能利用养分策略,从土壤养分变化、根系养分吸收转运、叶片生理过程等方面的整合研究思路,探讨氮沉降影响我国草地群落物种多样性的生物学机制,为我国草地生态系统的科学管理提供理论依据。     

大气CO2浓度升高对植物根系的影响

植物长期生长在CO2浓度不断升高的环境中,其结构和功能都将受到影响,这种影响不仅表现在植物的地上部分,同时也表现在植物的地下部分(根系),尤其是细根的长度、直径、产量、周转以及根与枝的分配模式等方面。植物根系结构和功能的改变影响植物地上部分和生态系统物质循环中的碳动态及土壤中碳库的变化。目前有关大气CO2浓度升高对根系动态影响的研究报道主要包括大气CO2浓度升高对根系结构(直径、分枝、长度、数量等)和根系生理(周转率、产量、碳分配模式等)的影响2个方面。目前,该领域研究还存在一些不足,例如在CO2浓度升高条件下,对植物根系内部的调控机制,以及由其引起的物质循环和能量流动的动态变化的了解较少;至今没有令人信服的证据说明大气CO2浓度升高使根系周转升高还是降低。今后应加强研究在CO2浓度升高条件下根系的周转变化和光合产物分配模式变化,CO2浓度升高和外界环境因素的共同作用对根系的影响,以及采用不同研究方法和研究对象在不同立地条件下开展升高CO2浓度对根系影响的对比研究等。     

The effect of elevated CO2 concentration and nutrient supply on carbon-based plant secondary metabolites in Pinus sylvestris L.

This study investigated changes in carbon-based plant secondary metabolite concentrations in the needles of Pinus sylvestris saplings, in response to long-term elevation of atmospheric CO₂, at two rates of nutrient supply. Experimental trees were grown for 3 years in eight open-top chambers (OTCs), four of which were maintained at ambient (∼350 μmol mol⁻¹) and four at elevated (700 μmol mol⁻¹) CO₂ concentrations, plus four open air control plots. Within each of these treatments, plants received either high (7.0 g N m⁻² year⁻¹ added) or low (no nutrients added) rates of nutrient supply for two years. Needles from lateral branches were analysed chemically for concentrations of condensed tannins and monoterpenes. Biochemical determinations of cellulase digestibility and protein precipitating capacity of their phenolic extracts were made because of their potential of importance in ecological interactions between pine and other organisms including herbivores and decomposers. Elevated CO₂ concentration caused an increase (P<0.05) in dry mass per needle, tree height and the concentration of the monoterpene α-pinene, but there were no direct effects of CO₂ concentration on any of the other chemical measurements made. High nutrient availability increased cellulase digestibility of pine needles. There was a significant negative effect of the OTCs on protein precipitating capacity of the needle extracts in comparison to the open-air controls. Results suggest that predicted changes in atmospheric CO₂ concentration will be insufficient to produce large changes in the concentration of condensed tannins and monoterpenes in Scots pine. Processes which are influenced by these compounds, such as decomposition and herbivore food selection, along with their effects on ecosystem functioning, are therefore unlikely to be directly affected through changes in these secondary metabolites. Received: 20 October 1997 / Accepted: 28 February 1998     

The stimulatory effect of bilirubin on phagocytic activity

The effect of an i.p. application of bilirubin on phagocytic activity and the arrangement of the microtubule system of murine peritoneal macrophages were examined. A significant increase of phagocytic activity was observed 3 h after bilirubin treatment, and normal values were reached after 3 d. The increase of the extent of the microtubule system was comparable with that of enhanced phagocytosis but normal values were observed after 1 d.     

不同尺度上植物叶气孔导度对升高CO2的响应

植物叶气孔导度对大气CO2浓度升高的响应可表现在以下几个层面：在叶水平上,叶气孔导度和气孔密度下降;在植物个体水平上,单位叶面积蒸腾下降,植株的水分利用率升高;在生态系统水平上,蒸散降低,土壤泾流和土壤水分含量增加;在全球尺度上,扩大了温室气体的增温效应,同时也降低了全球降雨量增加的趋势。正是因为植物叶气孔导度的变化会影响全球水循环,所以它在全球变化中起着非常重要的作用。但目前的研究结果还不能外推到更大的尺度上去。     

Influence of rhizodeposition under elevated CO2 on plant nutrition and soil organic matter

Atmospheric CO₂ concentrations can influence ecosystem carbon storage through net primary production (NPP), soil carbon storage, or both. In assessing the potential for carbon storage in terrestrial ecosystems under elevated CO₂, both NPP and processing of soil organic matter (SOM), as well as the multiple links between them, must be examined. Within this context, both the quantity and quality of carbon flux from roots to soil are important, since roots produce specialized compounds that enhance nutrient acquisition (affecting NPP), and since the flux of organic compounds from roots to soil fuels soil microbial activity (affecting processing of SOM).From the perspective of root physiology, a technique is described which uses genetically engineered bacteria to detect the distribution and amount of flux of particular compounds from single roots to non-sterile soils. Other experiments from several labs are noted which explore effects of elevated CO₂ on root acid phosphatase, phosphomonoesterase, and citrate production, all associated with phosphorus nutrition. From a soil perspective, effects of elevated CO₂ on the processing of SOM developed under a C4 grassland but planted with C3 California grassland species were examined under low (unamended) and high (amended with 20 g m^–2 NPK) nutrients; measurements of soil atmosphere 13C combined with soil respiration rates show that during vegetative growth in February, elevated CO₂ decreased respiration of carbon from C4 SOM in high nutrient soils but not in unamended soils.This emphasis on the impacts of carbon loss from roots on both NPP and SOM processing will be essential to understanding terrestrial ecosystem carbon storage under changing atmospheric CO₂ concentrations.Abbreviations SOM soil organic matter - NPP net primary productivity - NEP net ecosystem productivity - PNPP p-nitrophenyl phosphate     

The mechanisms of plant photosynthetic acclimation to elevated CO2 concentration]

When measured at a same CO(2) concentration, net photosynthetic rate is often significantly lower in long-term high CO(2)-grown plants than the ambient CO(2)-grown ones. This phenomenon is termed photosynthetic acclimation or down-regulation. Although there have been many reports and reviews, the mechanism(s) of the photosynthetic acclimation is not very clear. Combining the work of the authors' group, this paper briefly reviews the progress in studies on the mechanism(s) of the photosynthetic acclimation to elevated CO(2). It is suggested that besides the possible effects of respiration enhancement and excessive photosynthate accumulation, RuBP carboxylation limitation and RuBP regeneration limitation are probably the main factors leading to the photosynthetic acclimation.     

The resprouting response of co‐occurring temperate woody plant and grass species to elevated [CO2]: An insight into woody plant encroachment of grasslands

The phenomenon of woody plant thickening in grasslands has been observed globally and is likely to have widespread ecological consequences. It has been proposed that woody plant thickening is driven in part by rising atmospheric [CO₂] enhancing the resprouting ability of woody plants relative to grasses so they respond more strongly after disturbances such as herbivory and fire. The aim of this study was to examine the CO₂ effect on the resprouting ability of 16 co‐occurring temperate woody plant and grass species (eight species from each growth form). Plants were grown in a controlled glasshouse experiment under ambient (400 ppm) and elevated [CO₂] (600 ppm) for 14 weeks after which their resprouting ability was measured. Root non‐structural carbohydrate (NSC_mass) and nitrogen (N_mass) storage was used as proxies to measure the resprouting ability of woody plants while for the grasses it was measured directly. We found that both the woody plants (22% on average; P = 0.003) and grasses (20% on average; P = 0.003) produced more biomass under elevated [CO₂]. Despite the woody plants not allocating additional carbon to belowground storage under elevated [CO₂], they had significantly greater root NSC_mass (23% on average; P = 0.007) due to increased root biomass production (8% on average; P = 0.007). In contrast, root N_mass of the woody plants did not differ between CO₂ treatments (P = 0.373). Surprisingly, the resprouting ability of the grasses did not significantly differ between the CO₂ treatments (P = 0.067). These results provide evidence that the differing resprouting response of woody plants and grasses under elevated [CO₂] may be contributing to woody plant encroachment of grasslands worldwide.     

A link between plant diversity, elevated CO2 and soil nitrate

Interactive effects of reductions in plant species diversity and increases in atmospheric CO₂ were investigated in a long-term study in nutrient-poor calcareous grassland. Throughout the experiment, soil nitrate was persistently increased at low plant species diversity, and CO₂ enrichment reduced soil [NO₃^-] at all levels of plant species diversity. In our study, soil [NO₃^-] was unrelated to root length density, microbial biomass N, community legume contents, and experimental plant communities differed only little in total N pools. However, potential nitrification revealed exactly the same treatment effects as soil [NO₃^-], providing circumstantial evidence that nitrification rates drove the observed changes in [NO₃^-]. One possible explanation for plant diversity effects on nitrification lies in spatial and temporal interspecific differences in plant N uptake, which would more often allow accumulation of NH₄⁺ in part of the soil profile at low diversity than in more species-rich plant communities. Consequently, nitrification rates and soil [NO₃^-] would increase. Elevated CO₂ increased soil water contents, which may have improved NO₃^- diffusion to the root surface thereby reducing soil [NO₃^-]. Higher soil moisture at elevated CO₂ might also reduce nitrification rates due to less aerobic conditions. The accordance of the diversity effect on soil [NO₃^-] with previous experiments suggests that increased soil [NO₃^-] at low species diversity is a fairly general phenomenon, although the mechanisms causing high [NO₃^-] may vary. In contrast, experimental evidence for effects of CO₂ enrichment on soil [NO₃^-] is ambiguous, and the antagonistic interaction of plant species reductions and elevated CO₂ we have observed is thus probably less universal.     

The effect of elevated CO2 on growth and competition in experimental phytoplankton communities

We report an experiment designed to identify the effect of elevated CO₂ on species of phytoplankton in a simple laboratory system. Major taxa of phytoplankton differ in their ability to take up CO₂, which might lead to predictable changes in the growth rate of species and thereby shifts in the composition of phytoplankton communities in response to rising CO₂. Six species of phytoplankton belonging to three major taxa (cyanobacteria, diatoms and chlorophytes) were cultured in atmospheres whose CO₂ concentration was gradually increased from ambient levels to 1000 parts per million over about 100 generations and then maintained for a further 200 generations at elevated CO₂. The experimental design allowed us to trace a predictive sequence, from physiological features to the growth response of species to elevated CO₂ in pure culture, from the growth response in pure culture to competitive ability in pairwise mixtures and from pairwise competitive ability to shifts in the relative abundance of species in the full community of all six species. CO₂ altered the dynamics of growth in a fashion consistent with known differences among major taxa in their ability to take up and use CO₂. This pure‐culture response was partly successful in predicting the outcome of competition in pairwise mixtures, especially the enhanced competitive ability of chlorophytes relative to cyanobacteria, although generally statistical support was weak. The competitive response in pairwise mixtures was a good predictor of changes in competitive ability in the full community. Hence, there is a potential for forging a logical chain of inferences for predicting how phytoplankton communities will respond to elevated CO₂. Clearly further extensive experiments will be required to validate this approach in the greater complexity found in diverse communities and environments of natural systems.     

Indices of plant N availability in an alpine grassland under elevated atmospheric CO2

The objective of this study was to estimate whether elevated atmospheric [CO₂] alters plant N availability in a native high-elevation grassland in the Swiss Alps using two integrative, relatively non-disruptive methods. Estimates based on seasonal net plant N uptake, and those based on the amounts of NH ₄ ⁺ -N plus NO ₃ ^– -N captured by ion exchange resin (IER) bags, did not differ in plots treated with ambient (355 L L^–1) and elevated (680 L L^–1) [CO₂] in either the second (1993) or third (1994) growing season under treatment with elevated [CO₂]. The results of this study suggest that the effects of rising atmospheric [CO₂] on plant N availability may be negligible in this grassland. The results also contrast the relatively large effects of elevated atmospheric [CO₂] (increases and decreases) reported for highly disturbed artificial systems.     

Effects of elevated CO2 on flowering phenology and nectar production of nectar plants important for butterflies of calcareous grasslands

Effects of elevated CO₂ on flowering phenology and nectar production were investigated in Trifolium pratense, Lotus corniculatus, Scabiosa columbaria, Centaurea jacea and Betonica officinalis, which are all important nectar plants for butterflies. In glasshouse experiments, juvenile plants were exposed to ambient (350 μl l⁻¹) and elevated (660 μl l⁻¹) CO₂ concentrations for 60–80 days. Elevated CO₂ significantly enhanced the development of flower buds in C. jacea. B. officinalis flowered earlier and L. corniculatus produced more flowers under elevated CO₂. In contrast, the number of flowers decreased in T. pratense. The amount of nectar per flower was not affected by elevated CO₂ in the tested legumes (T. pratense and L. corniculatus), but was significantly reduced (!) in the other forbs. Elevated CO₂ did not significantly affect nectar sugar concentration and composition. However, S. columbaria and C. jacea produced significantly less total sugar under elevated CO₂. The nectar amino acid concentration remained unaffected in all investigated plant species, whereas the total of amino acids produced per flower was reduced in all non-legumes. In addition, the amino acid composition changed significantly in all investigated species except for C. jacea. The observed effects are unexpected and are a potential threat to flower visitors such as most butterflies which have no alternative food resources to nectar. Changes in nectar production due to elevated CO₂ could also have generally detrimental effects on the interactions of flowers and their pollinators. Received: 12 September 1996 / Accepted: 9 September 1997     

The influence of plant species,fertilization and elevated CO2 on soil aggregate stability

We tested the effects of plant species, fertilization and elevated CO₂ on water-stable soil aggregation. Five annual grassland species and a plant community were grown in outdoor mesocosms for 4 years, with and without NPK fertilization, at ambient or elevated atmospheric CO₂ concentrations. Aggregate stability (resistance of aggregates to slaking) in the top 0.15 m of soil differed among plant species. However, the more diverse plant community did not enhance aggregate stability relative to most monocultures. Species differences in aggregate stability were positively correlated with soil active bacterial biomass, but did not correlate with root biomass or fungal length. Plant species did not affect aggregate stability lower in the soil profile (0.15–0.45 m), where soil biological activity is generally decreased. Elevated CO₂ and NPK fertilization altered many of the factors known to influence aggregation, but did not affect water-stable aggregation at either depth, in any of the plant treatments. These results suggest that global changes will alter soil structure primarily due to shifts in vegetation composition.     

Long-term effect of elevated CO2 on spatial differentiation of ribulose-1,5-bisphosphate carboxylase/oxygenase activity in Norway spruce canopy

Total in vitro activity of RuBPCO (ribulose-1,5-bisphosphate carboxylase/oxygenase) enzyme was assayed spectrophotometrically by the continuous measurement of 3-phosphoglycerate-dependent NADH oxidation in a coupled enzyme system. RuBPCO activities were found in the ranges 1.01–2.76 and 1.23–3.10 µmol(CO₂) m⁻² s^{− 1} in current Norway spruce needles growing in ambient (AC) and elevated (EC) CO₂ concentration, respectively. RuBPCO activity in AC needles from the upper layer (U) was 11–15 % higher compared to those from the middle (M) layer, and even 44–56 % higher compared to the lower (L) layer of spruce crown. Over the vegetation season, we observed a highly significant decrease of RuBPCO activity in the EC-U needles from 3.10 (July) to 1.60 (October) µmol(CO₂) m⁻² s⁻¹ as a consequence of downward feedback regulation. Moreover, this down-regulation was not caused by a non-specific decrease in total leaf nitrogen content.The work forms a part of the research supported by grants no. LN00A141 and OC E21.001 (Ministry of Education CR), VaV640/18/03 (Ministry of Environment CR), and by the Research Intention of ILE AS CR AV0Z6087904.