Changes in C storage by terrestrial ecosystems: How C-N interactions restrict responses to CO2 and temperature

A general model of ecosystem biogeochemistry was used to examine the responses of arctic tundra and temperate hardwood forests to a doubling of CO₂ concentration and to a 5°C increase in average growing season temperature. The amount of C stored in both ecosystems increased with both increased CO₂ and temperature. Under increased CO₂, the increase in C storage was due to increases in the C∶N ratio of both vegetation and soils. Under increased temperature, the increased C storage in the forest was due to a shift in N from soils (with low C∶N ratios) to vegetation (with high C∶N ratios). In the tundra, both a shift in N from soils to vegetation and an increase in C∶N ratios contributed to increased C storage under higher temperatures. Neither ecosystem sequestered N from external sources because the supply rate was low.     

土壤微生物生物量和呼吸强度对大气CO2浓度升高的响应

随着全球环境变化对陆地生态系统的影响逐渐成为公众和科学界关注的热点,CO2作为一种重要的温室气体受到格外重视.大气CO2浓度升高将直接影响陆地植物的光合作用[1].植物的光合产物约有20% ～ 50%被运送到地下,通过根系分泌及死亡输入土壤[2],因此大气CO2浓度升高将会间接影响土壤生态系统.长期以来,关于大气CO2浓度升高对农作物地上部分的研究较多,但关于大气CO2浓度升高对土壤特别是土壤微生物的影响的研究报道较少.     

气候变暖对陆地生态系统的影响

人类活动引起的温室效应导致全球气候变暖,气候变暖对全球生态环境的影响越来越受到人们的关注.作为人类赖以生存的环境主体,陆地生态系统对气候变暖将做出何种响应,更是人们关注的重点.植物物候的变化可以直观地反映某些气候变化,尤其是气候变暖.气候变暖影响植物的生长节律,进而引起植物与环境关系的改变及生态系统物质循环(如水和碳的循环)的改变.不同种类植物对气候变化的差异响应,会使植物间和动植物间的竞争与依赖关系发生深刻的变化,如北半球中高纬度地区植被生长季延长、植物提早开花、昆虫提早出现、鸟类提早产蛋以及冰川退缩、永冻土带融化、江河湖泊结冰推迟而融化提早等.本文主要从陆地生态系统的分布和演替两方面着眼,以植物和动物作为考察对象,系统论述了森林、草原、荒漠、湿地及农田等陆地生态系统在气候变暖背景下产生的变化,并从微观和宏观尺度上提出陆地生态系统变化的生态学机制,最后在技术和政策层面给出若干对策.     

Belowground nematode herbivores are resistant to elevated atmospheric CO2 concentrations in grassland ecosystems

Grasslands are considered to be one of the most sensitive ecosystems to rising atmospheric CO₂ concentrations, since, in addition to direct effects of elevated CO₂ on plant growth, indirect increases in water availability as an effect of elevated CO₂ may enhance primary production and alter plant community composition in these typically dry ecosystems. Moreover, grasslands support large populations of belowground herbivores that consume a major portion of plant biomass. The direct trophic link between herbivores and plants suggests that primary consumers may be particularly sensitive to rising CO₂ concentrations; however, the responses of belowground grassland herbivores have rarely been investigated. Here, we report the response of a range of herbivorous nematode populations to elevated CO₂ concentrations from three distinct grassland experiments. The three studies each involved 5–6 years of CO₂ fumigation, utilized natural or representative plant and soil communities, and were sampled at the end of the growing season. In the vast majority of cases, elevated CO₂ did not affect the abundance of nematode families; only two nematode families were significantly influenced by CO₂ enrichment (Anguinidae increased in one case and Hoplolaimidae decreased in another). Similarly, elevated CO₂ did not influence the total abundance, family richness, diversity or plant parasitic index of the herbivorous nematode community. These neutral responses to CO₂ enrichment occurred despite increased root production in all three experiments, suggesting a simultaneous antagonistic mechanism may have operated, possibly decreased root quality and/or changes in the soil environment. Whatever the mechanism, our findings suggest that herbivorous nematodes in grassland ecosystems are resistant to rising atmospheric CO₂ concentrations.     

Tillage frequency affects microbial metabolic activity and short-term changes in CO2 fluxes within 1 week in karst ecosystems

Journal of Soils and Sediments - Tillage disturbance can significantly affect soil microbial metabolic activity and CO2 fluxes. Nevertheless, the influence of different tillage frequencies on...     

Warming and elevated CO2 combine to increase microbial mineralisation of soil organic matter

The net annual exchange of carbon between the atmosphere and terrestrial ecosystems is of prime importance in determining the concentration of CO₂ ([CO₂]) in the atmosphere and consequently future climate. Carbon loss occurs primarily through soil respiration; it is known that respiration is sensitive to the global changes in [CO₂] and temperature, suggesting that the net carbon balance may change in the future. However, field manipulations of temperature and [CO₂] alter many important environmental factors so it is unclear how much of the observed alterations in soil respiration is due to changes of microbial function itself instead of changes to the physical and chemical environment. Here we focus on resolving the importance of changes in the microbial community in response to warming and elevated [CO₂] on carbon mineralisation, something not possible in field measurements. We took plant material and soil inocula from a long running experiment where native grassland had been exposed to both warming and elevated CO₂ and constructed a reciprocal transplant experiment. We found that the rate of decomposition (heterotrophic respiration) was strongly determined by the origin of the microbial community. The combined warming + elevated CO₂ treatment produced a soil community that gave respiration rates 30% higher when provided with shoot litter and 70% for root litter than elevated CO₂ treatment alone, with the treatment source of the litter being unimportant. Warming, especially in the presence of elevated CO₂, increased the size of the apparent labile carbon pool when either C₃ or C₄ litter was added. Thus, the metabolic activity of the soil community was affected by the combination of warming and elevated CO₂ such that it had an increased ability to mineralise added organic matter, regardless of its source. Therefore, soil C efflux may be substantially increased in a warmer, high CO₂ world. Current ecosystem models mostly drive heterotrophic respiration from plant litter quality, soil moisture and temperature but our findings suggest equal attention will need to be paid to capturing microbial processes if we are to accurately project the future C balance of terrestrial ecosystems and quantify the feedback effect on atmospheric concentrations of CO₂.     

Influence of soil properties on microbial C,N, and P in dry tropical ecosystems

Summary Relationships between soil physicochemical characteristics and soil microbial C, N, and P in Indian dry tropical ecosystems are discussed. The major ecosystem studies were on forest, savanna, cropped fields, and mine spoils. The highest microbial C, N, and P levels were recorded from the mixed forest and the lowest levels in 5-year-old mine spoil. Across the sites, microbial C ranged from 226 to 643 g g^-1, microbial N from 19 to 71 g g^-1, and microbial P from 9 to 28 g g^-1 soil. The proportion of soil organic C contained in the microbial biomass ranged from 2.2 to 5.0%. The microbial C: N ratio in these soils ranged from 7.4. to 13.1 and the microbial C: P ratio from 16.6 to 30.6. The concentrations of microbial C, N, and P were correlated with several soil properties and among themselves. The soil properties, in various linear combinations, explained 90–99% of the variability in the microbial nutrients. Grazing of the savanna had some effect on the level of microbial biomass, and as the mine spoil aged, the level of microbial C, N, and P also increased.     

Soil organic matter and the extracellular microbial matrix show contrasting responses to C and N availability

An emerging paradigm in soil science suggests microbes can perform ‘N mining’ from recalcitrant soil organic matter (SOM) in conditions of low N availability. However, this requires the production of extracellular structures rich in N (including enzymes and structural components) and thus defies stoichiometric expectation. We set out to extract newly synthesised peptides from the extracellular matrix in soil and compare the amino acid (AA) profiles, N incorporation and AA dynamics in response to labile inputs of contrasting C/N ratio. Glycerol was added both with and without an inorganic source of N (10% ¹⁵N labelled NH₄NO₃) to a soil already containing a large pool of refractory SOM and incubated for 10 days. The resulting total soil peptide (TSP) and extracellular pools were compared using colorimetric methods, gas chromatography, and isotope ratio mass spectrometry. N isotope compositions showed that the extracellular polymeric substance (EPS) contained a greater proportion of products formed de novo than did TSP, with hydrophobic EPS-AAs (leucine, isoleucine, phenylalanine, hydroxyproline and tyrosine) deriving substantially more N from the inorganic source provided. Quantitative comparison between extracts showed that the EPS contained greater relative proportions of alanine, glycine, proline, phenylalanine and tyrosine. The greatest increases in EPS-peptide and EPS-polysaccharide concentrations occurred at the highest C/N ratios. All EPS-AAs responded similarly to treatment whereas the responses of TSP were more complex. The results suggest that extracellular investment of N (as EPS peptides) is a microbial survival mechanism in conditions of low N/high C which, from an evolutionary perspective, must ultimately lead to the tendency for increased N returns to the microbial biomass. A conceptual model is proposed that describes the dynamics of the extracellular matrix in response to the C/N ratio of labile inputs.     

Interaction of acid rain and global changes: Effects on terrestrial and aquatic ecosystems

Both acid deposition and changes in the global atmosphere and climate affect terrestrial and aquatic ecosystems. In the atmosphere sulphate aerosols tend to increase haze, altering the global radiation balance. Increased nitrogen deposition to N-limited systems such as boreal forests results in increased growth and increased sequestration of atmospheric CO₂, slowing the increase in CO₂ levels in the atmosphere. Future reduction in S and N emissions may result in a trade-off -- better with respect to some effects of acid deposition and greenhouse warming, but worse with respect to others. Global warming may cause the incidence and severity of drought to increase. Mineralisation of N and oxidation of organic S compounds release pulses of SO₄, acid and Al to surface waters. Effects in lakes may include reduced deep water refugia for cold stenotherms, lower nutrient concentrations, and greater penetration of harmful UV radiation. Longer water renewal times cause declines in SO₄ and NO₃, due to increased in situ removal, but increases in base cations. The net result is increased internal alkalinity production. In areas characterised by cold winters, global warming may result in a major shift in hydrologic cycle, with snowmelt episodes occurring during the winter rather than the typical pattern of accumulation in the winter and melting in the spring. Increased storm frequency predicted for the future will cause increased frequency and severity of sea salt episodes in coastal regions. Predicting the interactions of regional and global environmental factors in the coming decades poses new challenges to scientists, managers and policy-makers.     

Consistent patterns of N distribution through soil profiles in diverse alpine and tundra ecosystems

We studied the vertical patterns of δ¹⁵nitrogen in total N and exchangeable NH₄⁺-N through soil profiles in diverse alpine and tundra ecosystems. Soil samples were analyzed from 11 sites located in three mountain areas: NW Caucasus (Russia), the Khibiny Mountains (NW Russia) and Abisko region (N Sweden). Despite differences in the profile patterns of organic matter, nitrogen accumulation and nitrogen availability, we found consistent patterns of ¹⁵N distribution through all studied soil profiles. The δ¹⁵N values of total N were in general about zero or positive in the surface horizon and increased with soil depth. In contrast with total N, the δ¹⁵N values of exchangeable NH₄⁺-N were in general about zero or negative in the surface horizons and decreased with soil depth. NH₄⁺-N was significantly ¹⁵N-depleted compared with total N in all mineral horizons, while in the surface organic horizons differences between isotopic composition of total N and NH₄⁺-N were mostly not significant. We do not know the exact mechanism responsible for ¹⁵N depletion of NH₄⁺-N with soil depth and further research needs to evaluate the contributions of natural processes (higher nitrification activity and biological immobilization of “lighter” NH₄⁺-N near the soil surface) or artifacts of methodological procedure (contribution of the ¹⁵N-enriched microbial N and dissolved organic N near the soil surface). Nevertheless, our finding gives a new possibility to interpret variability in foliar δ¹⁵N values of plant species with different rooting depth in alpine and tundra ecosystems, because plants with deeper root systems can probably consume “lighter” rather than “heavier” NH₄⁺-N.     

Different responses of soil CO2 and N2O emissions to simulated N deposition in forests with divergent N transformation characteristics

CO₂ and N₂O are important greenhouse gases that are related to soil mineralization–immobilization turnover and nitrification. To explore the responses of CO₂ and N₂O emissions to N deposition in forests with different N transformation characteristics, CO₂ and N₂O fluxes were measured in two NH₄NO₃ fertilized plots. One plot was in a temperate pine plantation in Heilongjiang Liangshui National Nature Reserve (LS) with slow and minimally coupled mineralization–immobilization turnover and a high nitrification rate. The other plot was in a subtropical bamboo forest in the Fujian Daiyun Mountain National Nature Reserve (DY) in China with rapid and coupled mineralization–immobilization turnover but a low nitrification rate. The results showed that CO₂ emissions in the DY with a high mineralization rate were greater than those in the LS. Cumulative CO₂ emissions were significantly enhanced by N addition in DY, but in LS, they were not affected. The mean N₂O fluxes in the control were 0.010 and 0.008 mg N m^?2 hr^?1 for LS and DY, respectively. High N addition stimulated N₂O emissions in both LS and DY, but the response ratio for N₂O flux in LS (8.6) was larger than that in DY (2.9). These results suggested that soils with rapid and coupled mineralization–immobilization turnover are beneficial to CO₂ emissions and their positive response to N deposition. A high nitrification rate contributed to high N₂O emissions and the sensitive response of N₂O emissions to N deposition.     

Removal of atmospheric methane by soil ecosystems and its controlling variables from microbial to global scales

<正>Methane (CH₄), a potent greenhouse gas, plays a pivotal role in the dynamics of climate change. While CH₄ emissions have been widely investigated,biological removal of CH₄ by upland soils has been less explored. Understanding the mechanisms and factors affecting CH₄ oxidation in soils is of paramount importance for devising successful mitigation strategies. This perspective paper discusses different types of aerobic methanotrophs and their act...     

大气CO2和O3升高对菜地土壤酶活性和微生物量的影响

利用OTC平台和青菜盆栽实验,探索[CO2]、[O3]或[CO2+O3]升高条件下,土壤理化性质、微生物量和土壤酶活性的变化,以期获得未来大气CO2或/和O3升高对土壤微生态系统的风险性。结果表明,[CO2]升高不同程度地提高了土壤的可溶性有机碳（DOC）、可溶性有机氮（DON）、总磷（TP）、总碳（TC）、铵态氮（AN）、硝态氮（NN）含量和含水量（SWC）,进而不同程度地提高了土壤微生物量碳（MBC）、微生物量氮（MBN）含量以及土壤蛋白酶（PRA）、蔗糖酶（SA）、脲酶（UA）、多酚氧化酶（POA）、酸性磷酸酶（APA）和中性磷酸酶（NPA）活性。相反,[O3]升高不同程度降低了土壤DOC、TP、TK、TC、TN、AN、NN、SWC、MBC和MBN含量,提高了MBC/MBN比值,在不同程度上降低了土壤PRA、SA、UA、POA、APA和NPA酶活性。而[CO2+O3]在一定程度上消减了[O3]对土壤微生物量和酶活性的抑制作用,也降低了[CO2]升高对土壤微生物量和酶活性的刺激效应。因此,土壤微生物量和土壤酶活性的变化可用于评价未来大气CO2或/和O3升高对菜地土壤微生态环境的影响。     

Rapid response of soil protozoa to elevated CO2

Short-term changes in bacterial and protozoan populations from the soil of plants grown under elevated atmospheric CO₂ were quantified. We grew Brassica nigra at either ambient or twice-ambient CO₂ levels within open-top chambers in the field for 4 weeks. Plant biomass, above- and belowground, was unaffected by elevated CO₂. Direct count bacterial density was unchanged under elevated CO₂. Flagellate density tended to increase, whereas amoebal density significantly declined under elevated CO₂. This change in protozoan community structure suggests trophic transfer of the elevated CO₂ fertilization effect through the soil food chain. Received: 20 August 1996     

Short-term CO2 emissions from planted soil subject to elevated CO2 and simulated precipitation

Carbon dioxide emissions from soils beneath canopies of two Mediterranean plants, Artemisia absinthium L. and Festuca pratensis Huds. cv. Demeter, were monitored over a 7-day period that included an artificial precipitation event of 4 cm. The experiments were conducted using 0.2 m³ soil microcosms inside greenhouses with CO₂ concentrations of either 360 or 500 μmol mol⁻¹. Carbon dioxide flux from the soil surface, as calculated using a diffusive transport model agreed well with CO₂ flux measurements made using a dynamic flow system. Soil CO₂ emissions did not differ significantly between the 360 and 500 μmol mol⁻¹ CO₂ treatments when soils were dry (volumetric soil moisture content ≤9%). A simulated precipitation event caused an immediate exhalation of CO₂ from soil, after which CO₂ emissions declined slightly and remained constant for approximately 36 h. CO₂ emissions from soil microcosms with F. pratensis plants growing in 500 μmol mol⁻¹ CO₂ then rose to levels that were significantly greater than CO₂ emissions from soils in the microcosms exposed to 360 μmol mol⁻¹ CO₂. For A. absinthium growing in 500 μmol mol⁻¹ CO₂, the rise in soil CO₂ emissions following the wetting event was not significantly greater than emissions from soils with A. absinthium growing under 360 μmol mol⁻¹ CO₂. A. absinthium above ground biomass increased by 46.1 ± 17.9% (mean ± S.E., n = 4, P ≤ 0.05). Above ground biomass did not significantly increase for F. pratensis (14.4 ± 6.5%, P ≥ 0.10). Root biomass, on the other hand, increased for both species; by 50.6 ± 17.9% (P ≤ 0.05) for A. absinthium and by 55.9 ± 12.7% (P ≤ 0.05) for F. pratensis. Our results demonstrate two events following precipitation onto dry soils, an immediate release of CO₂ followed by a gradual increase from enhanced biological activity The gradual increase was greater for the herbaceous ruderal perennial F. pratensis under elevated CO₂.     

Composition and function of microbial communities during the early decomposition stages of foliar litter exposed to elevated CO2 concentrations

Foliar litter derived from trees exposed to elevated CO₂ concentration decreases in quality relative to foliar litter exposed to ambient CO₂. This change in quality could in turn affect the composition and function of microbial communities associated with litter transformation and mineralization. We evaluated the effects of decomposing litter obtained from three tree species, Pinus koraiensis, P. sylvestriformis and Quercus mongolicus, on the structure and function of microbial communities by burying litter‐bags and sampling the litters at intervals over a 429‐day period. Foliar litter collected from trees exposed to elevated CO₂ had greater C and lignin contents, and lost the mass at a slower rate than that collected from trees growing in ambient CO₂, except for P. koraiensis litter. The CO₂ responses of enzyme activity in litter were species‐dependent. Molecular fingerprinting by means of denaturing gradient gel electrophoresis (DGGE) demonstrated that elevated CO₂ more strongly affected the fungal community than the bacterial community in decomposing litter. In samples of P. koraiensis litter that had been exposed to elevated CO₂, sequences belonging to Basidiomycota showed increases in relative intensity, which were accompanied by significant increases in enzyme activities. Our results showed that enzyme activities and micro‐decomposer composition have different, litter‐specific responses to changes in litter biochemistry, which indicate that exposure of trees to elevated CO₂ has varying effects on litter decomposition.     

Influence of temperature on CO2 evolution,microbial biomass C and metabolic quotient during the decomposition of two humic forest horizons

The decomposition of the litter layer and the humic mineral horizon from a beech forest site was studied at temperatures of 5, 12, and 22°C for both substrates and additionally at 32°C for beech litter. Weight losses, basal and substrate-induced CO₂ production, and the extractable biomass C were monitored periodically during a 2-year incubation period. Weight losses and microbial activity were controlled by substrate quality and temperature. No significant differences were found between 5 and 12°C in decomposition, biomass C, and the metabolic quotient in the humic mineral horizon. The decay of beech litter and the humic mineral horizon was highest at 22°C but was faster in the litter material by a factor of 2.9 on average. In the glucose-amended samples, the relationship among the CO₂-C fluxes was 1:1:2:3 at temperatures of 5, 12, 22, and 32°C in the litter layer, and 1: 2: 2.4 at 5, 12, and 22°C in the A horizon, respectively. The microbial activity in the humic mineral horizon was only 2–11% of that in the litter layer. The level of biomass C remained constant over 1 year and no significant differences were obtained from the 12 and 22°C treatments in the litter layer.     

Contributions from different microbial processes to N2O emission from soil under different moisture regimes

Nitrous oxide emissions from a sandy-loam textured soil wetted to matric potentials of either-1.0 or-0.1 kPa were determined in laboratory experiments in which the soil was incubated in air (control), air plus 10 Pa C₂H₂ (to inhibit nitrification), 100 kPa O₂ (to suppress denitrification), 10 kPa C₂H₂ (to inhibit N₂O reduction to N₂ in denitrification) or following autoclaving. The total N₂O production, consumption and net N₂O emission from the soils together with the contributions to N₂O emission from different processes of N₂O production were estimated. The rate of N₂O production was significantly greater in the wetter soil (282 pmol N₂O g^-1 soil h^-1) than in the drier soil (192 pmol N₂O g^-1 soil h^-1), but because N₂O consumption by denitrifiers was also greater in the wetter soil, the net N₂O emissions from the wetter and the drier soils did not differ significantly. Non-biological sources made no significant contribution to N₂O emission under either moisture regime and biological processes other than denitrification and nitrification made only a small contribution (1% of the total N₂O production) in the wetter soil. Denitrifying nitrifiers were the predominant source of N₂O emitted from the drier soil and other (non-nitrifying) denitrifiers were the predominant source of N₂O emitted from the wetter soil.     

Effects of different N sources on N nutrient and microbial distribution across soybean rhizosphere

Abstract

Nitrogen is the most important nutrient for plant growth. In the present study, investigations were carried out on the effects of sodium nitrate, ammonium sulfate, urea, and two types of controlled-release coated urea (LP-40 and LP-70) fertilizers on the NO₃ ^?-N, NH₄ ⁺-N concentrations, and microbial numbers as well as pH distribution across the rhizosphere of soybean (Glycine max L. Merrill, var. Heinong 35). The study was conducted on a typical black soil using a rhizobox system. The results showed that NO₃ ^?-N was the main source of nitrogen, which was deficient in the rhizosphere in the treatments of ammonium sulfate, urea, LP-40, and LP-70, but accumulated considerably in the sodium nitrate treatment. The NH₄ ⁺-N concentration slightly increased in the rhizosphere in the ammonium sulfate treatment, and decreased in the rhizosphere when the other four kinds of N fertilizers were supplied. In an the treatments, bacterial and fungal numbers were highest in the central compartment (C.C.) of the rhizoboxes where the soybean root system was confined, but the rhizosphere width estimated from the increase in the microbial abundance differed among different N fertilizers. The experimental results also indicated that the fungal composition in the C.C. was less diverse than in other parts of the rhizobox compartments, and that the majority of fungal groups was represented by Penicillium spp., suggesting that the microbial distribution across the soybean rhizosphere differed both quantitatively and qualitatively.     

Modelling the response of terrestrial ecosystems to acidification and desiccation scenarios

Changes in vegetation are often caused by changes in abiotic site factors, such as pH, nitrogen availability and soil moisture. It has been recognized that abiotic site factors are affected by atmospheric deposition and groundwater-table changes. In order to evaluate the effects of eutrophication, acidification and desiccation on site factors, the model SMART2 has been developed. For the Netherlands combinations of two acidification and two seepage scenarios (1990–2050) were evaluated with SMART2. The results are focused on pH, nitrogen availability and base saturation. Calculations were made for combinations of five vegetation structures on seven soil types and the five groundwater-table classes, using a 1 km² grid. Results showed that deposition reductions lead to a relatively fast improvement of the site factors, increase in pH and base saturation and decrease in N availability. Whereas a reduction in groundwater abstractions of 25% has little or no effect on the pH and N availability.