Effects of mycorrhizal colonization on biomass production and nitrogen fixation of black locust (Robinia pseudoacacia) seedlings grown under elevated atmospheric carbon dioxide

Interactive effects of elevated atmospheric CO₂ and arbuscular mycorrhizal (AM) fungi on biomass production and N₂ fixation were investigated using black locust ( Robinia pseudoacacia ). Seedlings were grown in growth chambers maintained at either 350 μmol mol⁻¹ or 710 μmol mol⁻¹ CO₂. Seedlings were inoculated with Rhizobium spp. and were grown with or without AM fungi. The ¹⁵N isotope dilution method was used to determine N source partitioning between N₂ fixation and inorganic fertilizer uptake. Elevated atmospheric CO₂ significantly increased the percentage of fine roots that were colonized by AM fungi. Mycorrhizal seedlings grown under elevated CO₂ had the greatest overall plant biomass production, nodulation, N and P content, and root N absorption. Additionally, elevated CO₂ levels enhanced nodule and root mass production, as well as N₂ fixation rates, of non- mycorrhizal seedlings. However, the relative response of biomass production to CO₂ enrichment was greater in non-mycorrhizal seedlings than in mycorrhizal seedlings. This study provides strong evidence that arbuscular mycorrhizal fungi play an important role in the extent to which plant nutrition of symbiotic N₂-fixing tree species is affected by enriched atmospheric CO₂.     

Effect of elevated atmospheric CO2 on mycorrhizal colonization, external mycorrhizal hyphal production and phosphorus inflow in Plantago lanceolata and Trifolium repens in association with the arbuscular mycorrhizal fungus Glomus mosseae

Plantago lanceolata and Trifolium repens were grown under ambient (400 μmol mol^–1) and elevated (650 μmol mol^–1) atmospheric CO₂ conditions. Plants were inoculated with the arbuscular mycorrhizal fungus Glomus mosseae and given a phosphorus supply in the form of bonemeal. Six sequential harvests were taken in order to determine whether the effect of elevated CO₂ on internal mycorrhizal colonization and external hyphal production was independent of the stimulatory effect of elevated CO₂ on plant growth. At a given time, elevated CO₂ increased the percentage of root length colonized (RLC), the total length of colonized root and the external mycorrhizal hyphal (EMH) density and decreased the ratio of EMH to total length of colonized root. When plant size was taken into account, the CO₂ effect on RLC and total length of colonized root was greatly reduced (and only apparent for early harvests in T. repens) and the effects on the EMH parameters disappeared. Root tissue P concentration was unchanged at elevated CO₂, but there was a decrease in shoot P at the later harvests. There was no direct effect of elevated CO₂ on P inflow for the earlier period (< 50 d) of the experiment. However, over the last period, there was a significant negative effect of elevated CO₂ on P inflow for both species, independent of plant size. It is concluded that elevated CO₂ had no direct effect on mycorrhizal colonization or external hyphal production, and that any observed effects on a time basis were due to faster growing plants at elevated CO₂. However, for older plants, elevated CO₂ had a direct negative effect on P inflow. This decrease in P inflow coincides with the observed decrease in shoot P concentration. This is discussed in terms of downregulation of photosynthesis often seen in elevated CO₂ grown plants, and the potential for mycorrhizas (via external hyphal turnover) to alleviate the phenomenon. The direction for future research is highlighted, especially in relation to carbon flow to and storage in the soil.     

Effect of enhanced atmospheric CO2 on mycorrhizal colonization by Glomus mosseae in Plantago lanceolata and Trifolium repens

Plantago lanceolata L. and Trifolium repens L. were grown for 16 wk in ambient (360 μmol mol⁻¹) and elevated (610 μmol mol⁻¹) atmospheric CO₂. Plants were inoculated with the arbuscular mycorrhizal (AM) fungus Glomus mosseae (Nicol. & Gerd.) Gerdemann & Trappe and given a phosphorus supply in the form of bonemeal, which would not be immediately available to the plants. Seven sequential harvests were taken to determine whether the effect of elevated CO₂ on mycorrhizal colonization was independent of the effect of CO₂ on plant growth. Plant growth analysis showed that both species grew faster in elevated CO₂ and that P. lanceolata had increased carbon allocation towards the roots. Elevated CO₂ did not affect the percentage of root length colonized (RLC); although total colonized root length was greater, when plant size was taken into account this effect disappeared. This finding was also true for root length colonized by arbuscules. No CO₂ effect was found on hyphal density (colonization intensity) in roots. The P content of plants was increased at elevated CO₂, although both shoot and root tissue P concentration were unchanged. This was again as a result of bigger plants at elevated CO₂. Phosphorus inflow was unaffected by CO₂ concentrations. It is concluded that there is no direct permanent effect of elevated CO₂ on mycorrhizal functioning, as internal mycorrhizal development and the mycorrhizal P uptake mechanism are unaffected. The importance of sequential harvests in experiments is discussed. The direction for future research is highlighted, especially in relation to C storage in the soil.     

Effect of elevated CO2 on carbon partitioning and exudate release from Plantago lanceolata seedlings

Plantago lanceolata L. seedlings were grown in sand microcosm units over a 43‐day experimental period under two CO₂ regimes (800 or 400 µmol mol⁻¹) to investigate the effect of elevated atmospheric CO₂ concentration on carbon partitioning and exudate release. Total organic carbon (TOC) content of the collected exudate material was measured throughout the experimental period. After 42 days growth the seedlings were labelled with [¹⁴C]‐CO₂ and the fate of the label within the plant and its release by the roots monitored. Elevated CO₂ significantly (P ≤ 0.001) enhanced shoot, root and total dry matter production although the R:S ratio was unaltered, suggesting no alteration in gross carbon partitioning. The cumulative release of TOC (in mg C) over 0‐42 days was unaltered by CO₂ treatment however, when expressed as a percentage of net assimilated C, ambient‐grown plants released a significantly (P≤ 0.001) higher percentage from their roots compared to elevated CO₂‐grown plants (i.e. 8 vs 3%). The distribution of ¹⁴C‐label was markedly altered by CO₂ treatment with significantly (P≤ 0.001) greater per cent label partitioned to the roots under elevated CO₂. This indicates increased partitioning of recent assimilate below‐ground under elevated CO₂ treatment although there was no significant difference in the percentage of ¹⁴C‐label released by the roots. Comparison of plant C budgets based on ¹⁴C‐pulse‐chase methodology and TOC measurements is discussed.     

Plant and fungal responses to elevated atmospheric carbon dioxide in mycorrhizal seedlings of Pinus sylvestris

The effects of elevated CO₂ concentration upon the mycorrhizal relationships of Scots pine (Pinus sylvestris) seedlings were investigated. Plants were grown for 4 months with their shoots exposed to ambient (C_AMB=360 μl l⁻¹) or elevated (C_ELEV=700 μl l⁻¹) CO₂ environments while their root systems, either colonised by the mycorrhizal fungi Paxillus involutus or Suillus bovinus, or left in the non-mycorrhizal condition, were maintained in sealed dishes. In one series of these plants the effects of C_ELEV upon the extent of mycorrhizal development and upon their growth and nutrition were determined, while another series were transferred from the dishes after 1 month, to transparent observation chambers before being returned to the two CO₂ environments. In these chambers, the effects of C_ELEV upon development of the external mycelial systems of the two mycorrhizal fungi was determined by measuring the advance of the hyphal fronts of the mycorrhizal fungi across non-sterile peat from the colonised plants. The dry mass and number of mycorrhizal tips were significantly higher in C_ELEV than in the C_AMB condition in plants colonised by both fungi in the dishes. Yields of whole plants and of shoots were higher in the C_ELEV treatment whether or not they were grown in the mycorrhizal condition, but the greater yields were not associated in these sealed systems with enhanced nutrient gain. The dry mass of non-mycorrhizal plants was greater than that of those colonised by mycorrhizal fungi under elevated CO₂. This is thought to be attributable to the energetic cost of production of the larger mycorrhizal systems in this treatment. The extent of development of the mycorrhizal mycelial systems of both fungi was greatly increased in C_ELEV relative to that in C_AMB environments. It is hypothesised that increased allocation of carbon to mycorrhizal root systems and their associated mycelia would provide the potential for enhancement of nutrient acquisition in open systems of greater fertility.     

Effects of elevated atmospheric carbon dioxide on gas exchange and growth of white clover

Effects of rising atmospheric CO₂ concentrations on gas exchange, growth and productivity were investigated on an important grassland species, Trifolium repens L. cv. Blanca. Pure stands of this species were cultivated over an entire growing season in small acrylic greenhouses with an artificial atmosphere of ±367 or ±620 ppm CO₂, respectively. Effects on growth and development were examined in a functional growth analysis, while consequences for gas exchange were determined by photosynthesis and transpiration measurements on canopy level. The stands were regularly clipped for production assessment. Canopies grown at high CO₂ levels showed an average increase in productivity of almost 75%. Growth analysis indicated development of a larger foliage area as the major cause, particularly in the first days of regrowth after cutting. The growth advantage that began in this stage was maintained or bettered during the following weeks. The difference between gas exchange measurements expressed per unit leaf area and per unit ground area suggested that changes in net photosynthesis and respiration did not contribute to the increase in total yield. Transpiration declined under high CO₂ if expressed on a leaf area basis but total canopy transpiration was at least as large as in ambient CO₂ due to the larger leaf area. Water-use efficiency calculations on the summer data indicated a 35% improvement with a doubling of CO₂ concentration.     

Root production and mortality under elevated atmospheric carbon dioxide

An essential component of an understanding of carbon flux is the quantification of movement through the root carbon pool. Although estimates have been made using radiocarbon, the use of minirhizotrons provides a direct measurement of rates of root birth and death. We have measured root demographic parameters under a semi-natural grassland and for wheat. The grassland was studied along a natural altitudinal gradient in northern England, and similar turf from the site was grown in elevated CO₂ in solardomes. Root biomass was enhanced under elevated CO₂. Root birth and death rates were both increased to a similar extent in elevated CO₂, so that the throughput of carbon was greater than in ambient CO₂, but root half-lives were shorter under elevated CO₂ only under a Juncus/Nardus sward on a peaty gley soil, and not under a Festuca turf on a brown earth soil. In a separate experiment, wheat also responded to elevated CO₂ by increased root production, and there was a marked shift towards surface rooting: root development at a depth of 80–85 cm was both reduced and delayed. In conjunction with published results for trees, these data suggest that the impact of elevated CO₂ will be system-dependent, affecting the spatio-temporal pattern of root growth in some ecosystems and the rate of turnover in others. Turrnover is also sensitive to temperature, soil fertility and other environmental variables, all of which are likely to change in tandem with atmospheric CO₂ concentrations. Differences in turnover and time and location of rhizodeposition may have a large effect on rates of carbon cycling.     

Responses of soil biota to elevated atmospheric carbon dioxide

Increasing concentrations of atmospheric CO₂ could have dramatic effects upon terrestrial ecosystems including changes in ecosystem structure, nutrient cycling rates, net primary production, C source-sink relationships and successional patterns. All of these potential changes will be constrained to some degree by below ground processes and mediated by responses of soil biota to indirect effects of CO₂ enrichment. A review of our current state of knowledge regarding responses of soil biota is presented, covering responses of mycorrhizae, N-fixing bacteria and actinomycetes, soil microbiota, plant pathogens, and soil fauna. Emphasis will be placed on consequences to biota of increasing C input through the rhizosphere and resulting feedbacks to above ground systems. Rising CO₂ may also result in altered nutrient concentrations of plant litter, potentially changing decomposition rates through indirect effects upon decomposer communities. Thus, this review will also cover current information on decomposition of litter produced at elevated CO₂. Summary Predictably, the responses of soil biota to CO₂ enrichment and the degree of experimental emphasis on them increase with proximity to, and intimacy with, roots. Symbiotic associations are all stimulated to some degree. Total plant mycorrhization increases with elevated CO₂. VAM fungi increase proportionately with fine root length/mass increase. ECM fungi, however, exhibit greater colonization per unit root length/mass at elevated CO₂ than at current atmospheric levels. Total N-fixation per plant increases in all species examined, although the mechanisms of increase, as well as the eventual benefit to the host relative to N uptake may vary. Microbial responses are unclear. The assumption that changes in root exudation will drive increased mineralization and facilitate nutrient uptake should be examined experimentally, in light of recent models. Microbial results to date suggest that metabolic activity (measured as changes in process rates) is stimulated by root C input, rather than population size (measured by cell or colony counts). Insufficient evidence exists to predict responses of either soil-borne plant pathogens or soil fauna (i.e., food web responses). These are areas requiring attention, the first for its potential to limit ecosystem production through disease and the second because of its importance to nutrient cycling processes. Preliminary data on foliar litter decomposition suggests that neither nutrient ratios nor decomposition rates will be affected by rising CO₂. This is another important area that may be better understood as the number of longer term studies with more realistic CO₂ exposures increase. Evidence continues to mount that C fixation increases with CO₂ enrichment and that the bulk of this C enters the belowground component of ecosystems. The global fate and effects of this additional C may affect all hierarchical levels, from organisms to ecosystems, and will be largely determined by responses of soil biota.     

Response of plant roots to elevated atmospheric carbon dioxide

Plant root response to atmospheric CO₂ enrichment can be great. Results from this controlled environment investigation demonstrate substantial effects on root system architecture, micromorphology and physiology. The most pronounced effects were an increase in root length (110%) and root dry weight (143%). Root diameter, stele diameter, cortex width, root/shoot and root weight ratios all increased; root numbers did not increase. The long-term implications for belowground processes could be enormous.     

Impacts of elevated atmospheric ozone on peatland below-ground DOC characteristics

Rising concentrations of tropospheric ozone are having detrimental impacts on the growth of crop and forest species and some studies have reported inhibition of the allocation of carbon below ground. The effects of ozone on peatland ecosystems have received relatively little attention, despite their importance within the global carbon cycle. During this study, cores from a Welsh minerotrophic fen and ombrotrophic bog were exposed to four ambient/elevated ozone concentration regimes representing current and predicted 2050 profiles. A large and significant reduction in the concentration of porewater dissolved organic carbon (DOC) was recorded in the fen cores exposed to the elevated ozone concentrations (up to ?55%), with a concurrent shift to a higher molecular weight of the remaining soil carbon. No effects of ozone on DOC concentrations or characteristics were recorded for the bog cores. The data suggest higher ozone sensitivity of plants growing in the fen-type peatland, that the impacts on the vegetation may affect soil carbon characteristics through a reduction in root exudates and that there may have been a shift in the source of substrate DOC for microbial consumption from vegetation exudates to native soil carbon. There may also have been a direct effect of ozone molecules reacting with soil organic matter after being transported into the soil through the aerenchyma tissue of the overlying vegetation. These qualitative changes in the soil carbon in response to elevated ozone may have important implications for carbon cycling in peatland ecosystems, and therefore climate change.     

大气二氧化碳浓度升高对植物-昆虫相互关系的影响

综述了CO2浓度升高对植物与昆虫相互关系影响的研究结果。大量研究表明,高浓度CO2对植物生理生化活动有显著的影响,植物营养物质的变化对植食性昆虫亦产生不同程度的影响,高浓度CO2条件对咀嚼式口器昆虫的取食、生长发育和生殖有不同程度的不良影响,昆虫为了获得足够的氮素营养而增加取食强度和时间,从而更易于受到天敌的攻击,这些昆虫的生长率、繁殖和生存率有下降的趋势;而对刺吸韧皮部汁液的昆虫来说,多引起种群数量增加或无显著影响。并对研究中存在的问题进行了分析,提出了今后研究的方向。     

Effects of elevated carbon dioxide on three montane grass species: I. Growth and dry matter partitioning

Upland grasslands are a major component of natural vegetationwithin the UK. Such grasslands support slow growing relativelystable plant communities. The response of native montane grassspecies to elevated atmospheric carbon dioxide concentrationshas received little attention to date. Of such studies, mosthave only focused on short-term (days to weeks) responses, oftenunder favourable controlled environment conditions. In thisstudy Agrostis caplllaris L.⁵, Festuca vivipara L. and Poa alpinaL. were grown under semi-natural conditions in outdoor open-topchambers at either ambient (340µmol mol^–1) or elevated(680µmol mol^–1) concentrations of atmospheric carbondioxide (CO₂ for periods from 79 to 189 d, with a nutrient availabilitysimilar to that of montane Agrostis-Fescue grassland in Snowdonia,N. Wales. Whole plant dry weight was increased for A. capillarisand P. alpina, but decreased for F. vivipara, at elevated CO₂.Major components of relative growth rate (RGR) contributingto this change at elevated CO₂ were transient changes in specificleaf area (SLA) and leaf area ratio (LAR). Despite changes ingrowth rate at 680 µmol mol^–1 CO₂, partitioningof dry weight between shoot and root in plants of A. capillarisand P. alpina was unaltered. There was a significant decreasein shoot relative to root growth at elevated CO₂ in F. viviparawhich also showed marked discoloration of the leaves and increasedsenescence of the foliage. Key words: Allometry, growth analysis, elevated CO₂, grasses     

The effect of elevated atmospheric carbon dioxide levels on soil bacterial communities

The effect of elevated carbon dioxide levels on total bacterial communities was studied in a series of controlled and replicated model terrestrial ecosystems over a period of 38 weeks. The bacterial community was profiled using Denaturing Gradient Gel Electrophoresis (DGGE) analysis of bacterial 16S rRNA gene fragments amplified by the Polymerase Chain Reaction from DNA extracted directly from soil. Bacterial community DGGE profiles provided three major findings: (i) there was a high degree of profile similarity after ≈ 12 weeks (one plant generation); (ii) whilst overall DGGE profile was maintained over the 38 weeks (three plant generations), the banding patterns became more diverse with time; (iii) DGGE data provided no evidence for a shift in bacterial community structure resulting from exposure of the ecosystem to an increased atmospheric CO₂ level.     

The effects of increased atmospheric carbon dioxide and temperature on carbon partitioning, source-sink relations and respiration

Abstract. Herbaceous C₃ plants grown in elevated CO₂ show increases in carbon assimilation and carbohydrate accumulation (particularly starch) within source leaves. Although changes in the partitioning of biomass between root and shoot occur, the proportion of this extra assimilate made available for sink growth is not known. Root:shoot ratios tend to increase for CO₂-enriched herbaceous plants and decrease for CO₂-enriched trees. Root:shoot ratios for cereals tend to remain constant. In contrast, elevated temperatures decrease carbohydrate accumulation within source and sink regions of a plant and decrease root:shoot ratios. Allometric analysis of at least two species showing changes in root: shoot ratios due to elevated CO₂ show no alteration in the whole-plant partitioning of biomass. Little information is available for interactions between temperature and CO₂. Cold-adapted plants show little response to elevated levels of CO₂, with some species showing a decline in biomass accumulation. In general though, increasing temperature will increase sucrose synthesis, transport and utilization for CO₂-enriched plants and decrease carbohydrate accumulation within the leaf. Literature reports are discussed in relation to the hypothesis that sucrose is a major factor in the control of plant carbon partitioning. A model is presented in support.     

大气CO2浓度升高对花蜜及传粉昆虫的影响

植物依赖昆虫传授花粉,昆虫从植物获得花粉和花蜜作为食物,两者在漫长的进化过程中形成了密切的互惠共生关系.大量研究表明,CO2浓度升高对植物花蜜的产量和组成有显著的影响.CO2浓度增加后,有花植物花蜜的产量和组分在不同物种之间的变化差异很大,即使是种内不同基因型植株的花蜜对CO2浓度增加的反应也有所不同.大部分种类花蜜的...