Rhizodeposition under ambient and elevated CO2 levels

As global CO₂ levels rise, can soils store more carbon and so buffer atmospheric CO₂ levels? Answering this question requires a knowledge of the rates of C inputs to soil and of CO₂ outputs via decomposition. Below-ground inputs from roots are a major component of the C flow into soils but are still poorly understood. In this article, new techniques for measuring rhizodeposition are reviewed and discussed and the need for cross-comparisons between methods is identified. One component of rhizodeposition, root exudation, is examined in more detail and evidence is presented which suggests that current estimates of exudate flow into soils are incorrect. A mechanistic mathematical model is used to explore how exudate flows might change under elevated CO₂.     

桃蚜取食行为对大气CO_2浓度升高的响应

【目的】近年来随着人类活动的增加,温室气体尤其是大气CO_2浓度升高造成的虫害爆发已成为国际上关注的焦点,因此,研究拟南芥Arabidopsis thaliana上桃蚜取食行为的变化对大气CO_2浓度升高的响应意义重大。【方法】本研究以拟南芥和绿色桃蚜Myzus persicae为研究对象,利用野生型拟南芥Col-0,茉莉酸途径信号传导缺失突变体(jar1)、水杨酸途径信号传导缺失突变体(npr1)、乙烯途径信号传导缺失突变体(ein2-5)为材料,以大气CO_2浓度升高为影响因子,利用刺吸式电位仪(EPG)记录了桃蚜在不同处理的拟南芥上的取食波形。【结果】研究结果发现:CO_2浓度升高缩短了Col-0和jar1植株上蚜虫首次刺探时间和首次到达韧皮部的时间,却延长了npr1和ein2-5上蚜虫首次到达韧皮部的时间,降低了jar1植株上蚜虫总的刺探时间且增加了其总的取食韧皮部时间,但没有改变其它基因型植株上蚜虫总的刺探和取食时间;同时增加了野生型植株上蚜虫的刺探频率,却没有影响其它基因型植株上的刺探频率。【结论】CO_2浓度升高降低了野生型植株和jar1植株抗性,有利于蚜虫到达韧皮部;却增加了npr1和ein2-5上的植物抗性,从而不利于蚜虫到达韧皮部。     

Proteomic response of rice seedling leaves to elevated CO2 levels

Previous investigations of plant responses to higher CO 2 levels were mostly based on physiological measurements and biochemical assays. In this study, a proteomic approach was employed to investigate plant response to higher CO 2 levels using rice as a model. Ten-day-old seedlings were progressively exposed to 760 ppm, 1140 ppm, and 1520 ppm CO 2 concentrations for 24 h each. The net photosynthesis rate ( P n), stomatal conductance ( G s), transpiration rate ( E), and intercellular to ambient CO 2 concentration ratio ( C i/ C a) were measured. P n, G s, and E showed a maximum increase at 1140 ppm CO 2, but further exposure to 1520 ppm for 24 h resulted in down regulation of these. Proteins extracted from leaves were subjected to 2-DE analysis, and 57 spots showing differential expression patterns, as detected by profile analysis, were identified by MALDI-TOF/TOF-MS. Most of the proteins belonged to photosynthesis, carbon metabolism, and energy pathways. Several molecular chaperones and ascorbate peroxidase were also found to respond to higher CO 2 levels. Concomitant with the down regulation of P n and G s, the levels of enzymes of the regeneration phase of the Calvin cycle were decreased. Correlations between the protein profiles and the photosynthetic measurements at the three CO 2 levels were explored.     

The combined effects of elevated CO2 levels and UV-B radiation on growth characteristics ofElymus athericus (= E. pycnanathus)

Elymus athericus (Link) Kerguélen, a C₃ grass, was grown in a greenhouse experiment to determine the effect of enhanced atmospheric CO₂ and elevated UV-B radiation levels on plant growth. Plants were subjected to the following treatments; a) ambient CO₂-control UV-B, b) ambient CO₂-elevated UV-B, c) double CO₂-control UV-B, d) double CO₂-elevated UV-B. Elevated CO₂ concentrations stimulated plant growth, biomass production was 67% higher than at ambient CO₂. Elevated UV-B radiation had a negative effect on growth, biomass production was depressed by 31%. Enhanced CO₂ combined with elevated UV-B levels caused a biomass depression of 8% when compared with the control plants. UV-B induced growth depression can be modified by a growth stimulus caused by high CO₂ concentrations. Growth analysis has been performed and possible physiological mechanisms behind changing growth parameters are discussed.     

The effects of chamber pressurization on soil-surface CO2 flux and the implications for NEE measurements under elevated CO2

Soil and ecosystem trace gas fluxes are commonly measured using the dynamic chamber technique. Although the chamber pressure anomalies associated with this method are known to be a source of error, their effects have not been fully characterized. In this study, we use results from soil gas-exchange experiments and a soil CO₂ transport model to characterize the effects of chamber pressure on soil CO₂ efflux in an annual California grassland. For greater than ambient chamber pressures, experimental data show that soil-surface CO₂ flux decreases as a nonlinear function of increasing chamber pressure; this decrease is larger for drier soils. In dry soil, a gauge pressure of 0.5 Pa reduced the measured soil CO₂ efflux by roughly 70% relative to the control measurement at ambient pressure. Results from the soil CO₂ transport model show that pressurizing the flux chamber above ambient pressure effectively flushes CO₂ from the soil by generating a downward flow of air through the soil air-filled pore space. This advective flow of air reduces the CO₂ concentration gradient across the soil–atmosphere interface, resulting in a smaller diffusive flux into the chamber head space. Simulations also show that the reduction in diffusive flux is a function of chamber pressure, soil moisture, soil texture, the depth distribution of soil CO₂ generation, and chamber diameter. These results highlight the need for caution in the interpretation of dynamic chamber trace gas flux measurements. A portion of the frequently observed increase in net ecosystem carbon uptake under elevated CO₂ may be an artifact resulting from the impact of chamber pressurization on soil CO₂ efflux.     

The effects of elevated atmospheric CO2 and soil P placement on cotton root deployment

Root proliferation into nutrient rich zones is an important mechanism in the exploitation of soil nutrients by plants. No studies have examined atmospheric CO₂ effects on cotton (Gossypium hirsutum L.) root distribution as affected by localized phosphorus (P). Cotton plants were grown in a Troup sand (loamy, thermic Grossarenic Kandiudults) using 17.2-l containers placed in open top field chambers (OTC) under ambient (360 mol mol^–1) or enriched (720 mol mol^–1) atmospheric CO₂ concentrations for 40 days. Equivalent amounts of P were added (150 mg P per kg of soil) to 100, 50, 25, 12.5, and 6.25% of the total soil volume; control containers with no added P were also included. Under extremely low P (controls), cotton was unresponsive to CO₂ enrichment. In treatments with both fertilized and unfertilized soil volumes, root proliferation was greater in the unfertilized soil under elevated CO₂ conditions. Stimulation of root growth occurred in the P-fertilized soil fraction; the pattern of stimulation was similar under both CO₂ levels. Under ambient CO₂, cotton plant response was positive (shoot mass, and total root mass and length) when soil P was confined to relatively small proportions of the total soil volume (6.25 and 12.5%). However, elevated CO₂ grown plants tended to respond to P regardless of its distribution.     

The response of plants to elevated CO2

Summary Communities, consisting of six co-occurring, disturbed site annuals, were subjected to CO₂ unenriched (300 ppm) and to CO₂ enriched (450 and 600 ppm) atmospheres at different levels of light and nutrient availability. In general, total community production increased with CO₂ enrichment to 450 ppm, but a further increase in CO₂ to 600 ppm had little or no effect. The response of community production to CO₂ level was not affected by nutrient availability but was affected by light level.Of the six species, four display C₃ metabolism. The proportion of total community production contributed by these species increased as a result of CO₂ enrichment, and was dependent upon both light and nutrient availability. The relative success of some species, particularly in terms of reproduction (total seed biomass), was significantly altered by CO₂ concentration depending on the level of nutrients. There were not only changes in reproductive success (seed biomass) and shoot biomass but also changes in the proportion of biomass allocated to seed.These experiments demonstrate that CO₂ enrichment does affect annual plant communities both in terms of productivity and species composition and that the affect of CO₂ on such system may depend upon other resources such as light and nutrients.     

The response of plants to elevated CO2

Tree saplings, two groups of three species from each of two deciduous tree communities, were grown in competition at three CO₂ concentrations and two light levels. After one growing season, biomass was measured to assess the effect of CO₂ on community structure, and nitrogen and phosphorus concentrations were measured for leaves, stems, and roots of all trees. Gas-exchange measurements were made on the same species grown under the same CO₂ concentrations.Photosynthetic capacity (rate of photosynthesis at saturating CO₂ and light) tended to decline as CO₂ concentration increased, but differences were not statistically significant. Stomatal conductance declined significantly as CO₂ increased. Nitrogen and phosphorus concentrations generally declined as CO₂ increased, but there were some unexpected patterns in roots and stems. CO₂ concentration did not significantly affect the overall growth of either community after one season, but the relative biomass of each species changed in a complex way, depending on CO₂ light level, and community.     

The response of plants to elevated CO2

Summary Four coexisting annual plant species were grown in competition at three levels of CO₂ (300, 600, and 1,200 ppm) and two levels of soil moisture (moist and dry). Plant height was higher at high CO₂ concentrations for the three C₃ species but not for the C₄ species (Amaranthus retroflexus). Total community biomass increased with increasing CO₂ at both soil moisture levels. The contribution of each species to total community biomass was influenced by CO₂ concentration. The effects were especially pronounced for Polygonum pensylvanicum which contributed more to community production as CO₂ and soil moisture increased. Amaranthus behaved in exactly the reverse way; it did best under ambient CO₂ and dry soil moisture conditions. The results suggest that changes in competitive interactions and community structure will occur with the anticipated rise in global CO₂ concentration.     

The effects of elevated atmospheric CO2 on freshwater periphyton in a temperate stream

This study examines the effects of elevated CO₂ on the benthic biology of a temperate freshwater stream. We tested the hypotheses that elevated CO₂ would increase periphyton biomass, alter elemental composition, and change community composition by increasing the frequency of algal taxa most limited by CO₂ availability. Carbon dioxide was bubbled into reservoirs of stream water, increasing the ambient pCO₂ by approximately 1100 ppm. The CO₂-enriched water then flowed into artificial stream channels. Ceramic tiles were placed into the channels to allow for periphyton colonization. Dissolved inorganic carbon increased and pH decreased with added CO₂. Measurements of biological parameters including periphyton biomass, algal C:N:P ratios, and community composition suggest that the periphyton were unaffected by the changes in stream water chemistry. We infer that rising atmospheric CO₂ will impact stream water chemistry but that periphyton may not be the first to respond to these changes. Impacts to alkaline freshwater streams from elevated CO₂ initially may be due to changes to terrestrial inputs that affect microbial decomposition and grazer activity, rather than through increases in periphyton carbon fixation. However, environmental characteristics of freshwater systems vary considerably, and additional studies are needed for accurate predictive modeling and monitoring of the effects of increasing atmospheric CO₂ on freshwater streams.

     

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.     

Non-litter effects of elevated CO2 on forest floor microarthropod abundances

The arthropod assemblages of the litter and soil play significant roles in decomposition and nutrient cycling. At the FACE (Free-Air CO₂ Enrichment) site at the Duke Forest, we assessed the responses of the litter microarthropod assemblage to elevated CO₂ (200 ppm above ambient) in a loblolly pine plantation. Following the initiation of the elevated CO₂ treatment, a trend toward lower microarthropod abundance under elevated CO₂ emerged. After 18 months, the mean microarthropod abundance was 33% lower in the elevated treatment (P=0.04). The decline was evident in all microarthropod groups, but was significant only in the oribatid mites (P=0.04). Because these responses precede any changes in litter quality resulting from the CO₂ treatment, they may reflect plant-derived changes in the soil that are being conveyed into the litter layer.     

Evidence that elevated CO2 levels can indirectly increase rhizosphere denitrifier activity

We examined the influence of elevated CO2 concentration on denitrifier enzyme activity in wheat rhizoplanes by using controlled environments and solution culture techniques. Potential denitrification activity was from 3 to 24 times higher on roots that were grown under an elevated CO2 concentration of 1,000 micromoles of CO2 mol-1 than on roots grown under ambient levels of CO2. Nitrogen loss, as determined by a nitrogen mass balance, increased with elevated CO2 levels in the shoot environment and with a high NO3- concentration in the rooting zone. These results indicated that aerial CO2 concentration can play a role in rhizosphere denitrifier activity.     

Monoterpene levels in needles of Douglas fir exposed to elevated CO2 and temperature

Monoterpene levels in current year needles of Douglas fir ( Pseudotsuga menziesii (Mirb.) Franco) seedlings were measured at the end of 4 years of exposure to ambient or elevated CO₂ (+179 µmol mol⁻¹), and ambient or elevated temperature (+0.3.5^C). Eleven monoterpenes were identified and quantified using gas chromatography/flame ionization detector/mass spectroscopy, with eight of these compounds regularly occurring in all trees examined. Elevated CO₂ exposure significantly reduced the levels for four of the eight main compounds in needles. Total monoterpene production was reduced by 52% ( P  < 0.05). Elevated temperature also reduced monoterpene levels ( P  < 0.07). The combination of elevated temperature and elevated CO₂ resulted in a 64% reduction in total monoterpenes compared with needles on ambient temperature trees. Two-way anova showed no significant temperature-CO₂ interaction. It is hypothesized that seasonal reductions in needle monoterpene pools under elevated CO₂ and temperature conditions may be due to a combination of competing carbon sinks, including increased carbon flux through the roots.     

The effects of elevated CO2 on seed production and seedling recruitment in a sheep-grazed pasture

Seed production and seedling recruitment were measured over 2 years under ambient (360 ppm) and elevated (475 ppm) atmospheric CO₂ in a free air carbon dioxide enrichment (FACE) experiment, carried out in a sheep-grazed pasture on dry, sandy soil in New Zealand. In both years elevated CO₂ led to more dispersed seeds of the grasses Anthoxanthum odoratum, Lolium perenne and Poa pratensis, the legumes Trifolium repens and T. subterraneum and the herbs Hypochaeris radicata and Leontodon saxatilis. The increased seed dispersal in A. odoratum, H. radicata, Leontodon saxatilis and T. repens reflected both more inflorescences per unit area and more seeds per inflorescence under elevated CO₂. The increased seed dispersal in Lolium perenne, P. pratensis and T. subterraneum was due solely to more inflorescences per unit area. The number of seedlings that emerged and survived to at least 7 months of age was increased by elevated CO₂ for H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in both years and for A. odoratum and Lolium perenne in the first year. For species where increased seedling recruitment was noted, there was a significant positive correlation between seed production in summer and seedling emergence in the following autumn and winter, and sowing 200 extra seeds per species m^-2 resulted in more seedlings compared to unsown controls. Elevated CO₂ did not affect seedling survival in any species. There was no measurable effect of elevated CO₂ on canopy and soil surface conditions or soil moisture at the time of seedling emergence. The results suggest the dominant effect of elevated CO₂ on seedling recruitment in this pasture was an indirect one, reflecting effects on the number of seeds produced. The biomass of H. radicata, Leontodon saxatilis, T. repens and T. subterraneum in the above-ground vegetation was greater under elevated than ambient CO₂. However, the size of individual seedlings and mature plants of these four species was unaffected by elevated CO₂. The results indicate an important way elevated CO₂ influenced plant species composition in this pasture was through changes in the pattern of seedling recruitment.     

The interactive effects of elevated CO2 and O3 concentration on photosynthesis in spring wheat

This study investigated the interacting effects of carbon dioxide and ozone on photosynthetic physiology in the flag leaves of spring wheat (Triticum aestivum L. cv. Wembley), at three stages of development. Plants were exposed throughout their development to reciprocal combinations of two carbon dioxide and two ozone treatments: [CO₂] at 350 or 700 mol mol^–1, [O₃] at < 5 or 60 nmol mol^–1. Gas exchange analysis, coupled spectrophotometric assay for RuBisCO activity, and SDS-PAGE, were used to examine the relative importance of pollutant effects on i) stomatal conductance, ii) quantum yield, and iii) RuBisCO activity, activation, and concentration. Independently, both elevated [CO₂] and elevated [O₃] caused a loss of RuBisCO protein and V_cmax. In combination, elevated [CO₂] partially protected against the deleterious effects of ozone. It did this partly by reducing stomatal conductance, and thereby reducing the effective ozone dose. Elevated [O₃] caused stomatal closure largely via its effect on photoassimilation.     

The effects of elevated CO2 on root respiration rates of two Mojave Desert shrubs

Although desert ecosystems are predicted to be the most responsive to elevated CO₂, low nutrient availability may limit increases in productivity and cause plants in deserts to allocate more resources to root biomass or activity for increased nutrient acquisition. We measured root respiration of two Mojave Desert shrubs, Ambrosia dumosa and Larrea tridentata, grown under ambient (～375 ppm) and elevated (～517 ppm) CO₂ concentrations at the Nevada Desert FACE Facility (NDFF) over five growing seasons. In addition, we grew L. tridentata seedlings in a greenhouse with similar CO₂ treatments to determine responses of primary and lateral roots to an increase in CO₂. In both field and greenhouse studies, root respiration was not significantly affected by elevated CO₂. However, respiration of A. dumosa roots <1 month old was significantly greater than respiration of A. dumosa roots between 1 and 4 months old. For both shrub species, respiration rates of very fine (<1.0 mm diameter) roots were significantly greater than those of fine (1–2 mm diameter) roots, and root respiration decreased as soil water decreased. Because specific root length was not significantly affected by CO₂ and because field minirhizotron measurements of root production were not significantly different, we infer that root growth at the NDFF has not increased with elevated CO₂. Furthermore, other studies at the NDFF have shown increased nutrient availability under elevated CO₂, which reduces the need for roots to increase scavenging for nutrients. Thus, we conclude that A. dumosa and L. tridentata root systems have not increased in size or activity, and increased shoot production observed under elevated CO₂ for these species does not appear to be constrained by the plant's root growth or activity.     

Interactive effects of elevated CO2 and temperature on water transport inponderosa pine

Many studies report that water flux through trees declines in response to elevated CO₂, but this response may be modified by exposure to increased temperatures. To determine whether elevated CO₂ and temperature interact to affect hydraulic conductivity, we grew ponderosa pine seedlings for 24 wk in growth chambers with one of four atmospheric CO₂ concentrations (350, 550, 750, and 1100 ppm) and either a low (15°C nights, 25°C days) or high (20°C nights, 30°C days) temperature treatment. Vapor pressure deficits were also higher in the elevated temperature treatment. Seedling biomass increased with CO₂ concentration but was not affected by temperature. Root : shoot ratio was unaffected by CO₂ and temperature. Leaf : sapwood area ratio (A_L/A_S) declined in response to elevated temperature but was not influenced by CO₂. Larger tracheid diameters at elevated temperature caused an increase in xylem-specific hydraulic conductivity (K_S). The increase in K_S and decrease in A_L/A_S led to higher leaf-specific hydraulic conductivity (K_L) at elevated temperature. Stomatal conductance (g_S) was correlated with K_L across all treatments. Neither K_S, K_L, nor g_S were affected by elevated CO₂ concentrations. High K_L in response to elevated temperature may support increased transpiration or reduce the incidence of xylem cavitation in ponderosa pine in future, warmer climates.     

开放式空气CO2浓度增高对土壤线虫影响的研究现状与展望

大气CO2浓度增高会对生态系统产生一系列的影响,这些影响在某种程度上受到土壤动物区系的调节,本文通过论述大气CO2浓度增高对不同类型土壤中和不同生态系统中土壤线虫产生的影响,阐明了用土壤线虫作为指示生物来研究生态系统变化的意义,并提出了今后针对大气CO2浓度增高这一现象应着重围绕土壤线虫及土壤动物系优先开展的几方面研究,从而更好地指示整个生态系统的变化情况,为有效地管理农田生态系统提供依据。