The effects of elevated CO2 on symbiotic N2 fixation: a link between the carbon and nitrogen cycles in grassland ecosystems

The response of plants to elevated CO₂ is dependent on the availability of nutrients, especially nitrogen. It is generally accepted that an increase in the atmospheric CO₂ concentration increases the C:N ratio of plant residues and exudates. This promotes temporary N-immobilization which might, in turn, reduce the availability of soil nitrogen. In addition, both a CO₂ stimulated increase in plant growth (thus requiring more nitrogen) and an increased N demand for the decomposition of soil residues with a large C:N will result under elevated CO₂ in a larger N-sink of the whole grassland ecosystem. One way to maintain the balance between the C and N cycles in elevated CO₂ would be to increase N-import to the grassland ecosystem through symbiotic N₂ fixation. Whether this might happen in the context of temperate ecosystems is discussed, by assessing the following hypothesis: i) symbiotic N₂ fixation in legumes will be enhanced under elevated CO₂, ii) this enhancement of N₂ fixation will result in a larger N-input to the grassland ecosystem, and iii) a larger N-input will allow the sequestration of additional carbon, either above or below-ground, into the ecosystem. Data from long-term experiments with model grassland ecosystems, consisting of monocultures or mixtures of perennial ryegrass and white clover, grown under elevated CO₂ under free-air or field-like conditions, supports the first two hypothesis, since: i) both the percentage and the amount of fixed N increases in white clover grown under elevated CO₂, ii) the contribution of fixed N to the nitrogen nutrition of the mixed grass also increases in elevated CO₂. Concerning the third hypothesis, an increased nitrogen input to the grassland ecosystem from N₂ fixation usually promotes shoot growth (above-ground C storage) in elevated CO₂. However, the consequences of this larger N input under elevated CO₂ on the below-ground carbon fluxes are not fully understood. On one hand, the positive effect of elevated CO₂ on the quantity of plant residues might be overwhelming and lead to an increased long-term below-ground C storage; on the other hand, the enhancement of the decomposition process by the N-rich legume material might favour carbon turn-over and, hence, limit the storage of below-ground carbon.     

黄土高原人工苜蓿草地固碳效应评估

苜蓿草地作为黄土高原草地的重要组成部分,在黄土高原草地生态系统碳循环占有重要地位。同时由于苜蓿具有固氮和固碳的双重作用,使它在低碳农业的发展中具有重要意义。本文以一年生（建植初期）、三年生（盛产期）、五年生（稳产期）和十年生（衰败期）的人工苜蓿草地为研究对象,利用地上地下生物量和土壤碳贮量,定量分析了人工苜蓿草地不同发育阶段的固碳能力;结合种植面积,对黄土高原区人工苜蓿草地的总固碳效应进行了评价。研究结果表明：苜蓿草地地上生物固碳能力以三年生和五年生最强,并显著高于其他年限;地下生物固碳和土壤固碳量均以十年苜蓿草地最高,且显著高于其他年限。总体而言,人工苜蓿草地以土壤固碳为主,并集中在0~80cm土壤,各年限0-80cm土层土壤固碳量分别占到总固碳量的41.02%、39.43%、41.56%、39.59%;而地下生物固碳主要发生在0~20cm土层,各年限该层生物固碳量分别占到地下生物总固碳量的45.74%、 55.68%、53.12%和 43.28%。苜蓿草地在整个生命周期中总固碳量随生长年限逐年增长,但增长速度不明显,各年限单位面积总固碳量分别为12.69kg/m2、13.49 kg/m2、13.31 kg/m2和14.83 kg/m2。估算得到黄土高原的人工苜蓿草地年总固碳量为1.430?1011 Kg（143.0 Tg）,其中地上、地下、土壤分别为5.43Tg、1.15Tg和136.4Tg。本研究结果为正确分析区域碳循环和发展低碳大农业提供了重要理论参考。     

Phosphorus availability and elevated CO2 affect biological nitrogen fixation and nutrient fluxes in a clover-dominated sward

The response of biological nitrogen fixation (BNF) to elevated CO(2) was examined in white clover (Trifolium repens)-dominated swards under both high and low phosphorus availability. Mixed swards of clover and buffalo grass (Stenotaphrum secundatum) were grown for 15 months in 0.2 m2 sand-filled mesocosms under two CO2 treatments (ambient and twice ambient) and three nutrient treatments [no N, and either low or high P (5 or 134 kg P ha(-1)); the third nutrient treatment was supplied with high P and N (240 kg N ha(-1))]. Under ambient CO2, high P increased BNF from 410 to 900 kg ha(-1). Elevated CO2 further increased BNF to 1180 kg ha(-1) with high P, but there was no effect of CO2 on BNF with low P. Allocation of N belowground increased by approx. 50% under elevated CO2 irrespective of supplied P. The results suggest that where soil P availability is low, elevated CO2 will not increase BNF, and pasture quality could decrease because of a reduction in aboveground N.     

Mitigating the greenhouse gas balance of ruminant production systems through carbon sequestration in grasslands

Soil carbon sequestration (enhanced sinks) is the mechanism responsible for most of the greenhouse gas (GHG) mitigation potential in the agriculture sector. Carbon sequestration in grasslands can be determined directly by measuring changes in soil organic carbon (SOC) stocks and indirectly by measuring the net balance of C fluxes. A literature search shows that grassland C sequestration reaches on average 5 ± 30 g C/m2 per year according to inventories of SOC stocks and -231 and 77 g C/m2 per year for drained organic and mineral soils, respectively, according to C flux balance. Off-site C sequestration occurs whenever more manure C is produced by than returned to a grassland plot. The sum of on- and off-site C sequestration reaches 129, 98 and 71 g C/m2 per year for grazed, cut and mixed European grasslands on mineral soils, respectively, however with high uncertainty. A range of management practices reduce C losses and increase C sequestration: (i) avoiding soil tillage and the conversion of grasslands to arable use, (ii) moderately intensifying nutrient-poor permanent grasslands, (iii) using light grazing instead of heavy grazing, (iv) increasing the duration of grass leys; (v) converting grass leys to grass-legume mixtures or to permanent grasslands. With nine European sites, direct emissions of N2O from soil and of CH4 from enteric fermentation at grazing, expressed in CO2 equivalents, compensated 10% and 34% of the on-site grassland C sequestration, respectively. Digestion inside the barn of the harvested herbage leads to further emissions of CH4 and N2O by the production systems, which were estimated at 130 g CO2 equivalents/m2 per year. The net balance of on- and off-site C sequestration, CH4 and N2O emissions reached 38 g CO2 equivalents/m2 per year, indicating a non-significant net sink activity. This net balance was, however, negative for intensively managed cut sites indicating a source to the atmosphere. In conclusion, this review confirms that grassland C sequestration has a strong potential to partly mitigate the GHG balance of ruminant production systems. However, as soil C sequestration is both reversible and vulnerable to disturbance, biodiversity loss and climate change, CH4 and N2O emissions from the livestock sector need to be reduced and current SOC stocks preserved.     

Long-term effects of CO2 enrichment and temperature increase on the carbon balance of a temperate grass sward

Perennial ryegrass swards were grown in large containers ona soil, at two N fertilizer supplies and were exposed duringtwo years in highly ventilated plastic tunnels to elevated (700µl l^–1 [CO₂) or ambient atmospheric CO₂ concentrationat outdoor temperature and to a 3C increase in air temperaturein elevated CO₂. The irrigation was adjusted to obtain a soilwater deficit during summer. The daily net C assimilation wasincreased in elevated CO₂ by 29 and 36% at the low and highN supplies, respectively. Canopies grown in elevated CO₂ for14 to 27 months photosynthetized significantly less rapidly,in both elevated and normal CO₂ concentrations, than their counterpartsdeveloped in ambient CO₂ but the magnitude of this effect wassmall (–8% to –13%). Elevated CO₂ resulted in alarge increase in the fructan concentration in the pseudostemsand laminae (+46% and +189%, respectively). In elevated CO₂,the hexose and sucrose pool increased by 28% in the laminae,whereas it did not vary significantly in the pseudo-stems. A3C temperature increase in elevated CO₂ did not affect significantlythe average WSC concentrations in the pseudostems and laminae.The elevated CO₂ effects on the net C assimilation and on thenocturnal shoot respiration were greater in summer than in spring.On average, a 35% increase in the below-ground respiration wasmeasured in elevated CO₂. At the high N supply, a 3C increasein air temperature led to a decline in the below-ground respirationdue to a low soil moisture. The below-ground carbon storagewas increased by 32% and 96% in elevated CO₂ at the low andhigh N supplies, respectively, with no significant increasedtemperature effect. The role for the below-ground carbon storageof CO₂-induced changes in the root fraction of the grass andof temperature-induced changes in the moisture content of thesoil are discussed. Key words: Climate change, grassland, gas exchange, carbohydrates, carbon cycle     

Stimulation of Symbiotic N2 Fixation in Trifolium repens L. under Elevated Atmospheric pCO2 in a Grassland Ecosystem

Symbiotic N2 fixation is one of the main processes that introduces N into terrestrial ecosystems. As such, it may be crucial for the sequestration of the extra C available in a world of continuously increasing atmospheric CO2 partial pressure (pCO2). The effect of elevated pCO2 (60 Pa) on symbiotic N2 fixation (15N-isotope dilution method) was investigated using Free-Air-CO2-Enrichment technology over a period of 3 years. Trifolium repens was cultivated either alone or together with Lolium perenne (a nonfixing reference crop) in mixed swards. Two different N fertilization levels and defoliation frequencies were applied. The total N yield increased consistently and the percentage of plant N derived from symbiotic N2 fixation increased significantly in T. repens under elevated pCO2. All additionally assimilated N was derived from symbiotic N2 fixation, not from the soil. In the mixtures exposed to elevated pCO2, an increased amount of symbiotically fixed N (+7.8, 8.2, and 6.2 g m-2 a-1 in 1993, 1994, and 1995, respectively) was introduced into the system. Increased N2 fixation is a competitive advantage for T. repens in mixed swards with pasture grasses and may be a crucial factor in maintaining the C:N ratio in the ecosystem as a whole.     

Nitrous oxide emissions from grass swards during the eighth year of elevated atmospheric pCO2 (Swiss FACE)

Emissions of N₂O were measured during the growth season over a year from grass swards under ambient (360 μL L^?1) and elevated (600 μL L^?1) CO₂ partial pressures at the Free Air Carbon dioxide Enrichment (FACE) experiment, Eschikon, Switzerland. Measurements were made following high (56 g N m^?2 yr^?1) and low (14 g N m^?2 yr^?1) rates of fertilizer application, split over 5 re‐growth periods, to Lolium perenne, Trifolium repens and mixed Lolium/Trifolium swards. Elevated pCO₂ increased annual emissions of N₂O from the high fertilized Lolium and mixed Lolium/Trifolium swards resulting in increases in GWP (N₂O emissions) of 179 and 111 g CO₂ equivalents m^?2, respectively, compared with the GWP of ambient pCO₂ swards, but had no significant effect on annual emissions from Trifolium monoculture swards. The greater emissions from the high fertilized elevated pCO₂Lolium swards were attributed to greater below‐ground C allocation under elevated pCO₂ providing the energy for denitrification in the presence of excess mineral N. An annual emission of 959 mg N₂O‐N m^?2 yr^?1 (1.7% of fertilizer N applied) was measured from the high fertilized Lolium sward under elevated pCO₂. The magnitude of emissions varied throughout the year with 84% of the total emission from the elevated pCO₂Lolium swards measured during the first two re‐growths (April–June 2001). This was associated with higher rainfall and soil water contents at this time of year. Trends in emissions varied between the first two re‐growths (April–June 2001) and the third, fourth and fifth re‐growths (late June–October 2000), with available soil NO₃^? and rainfall explaining 70%, and soil water content explaining 72% of the variability in N₂O in these periods, respectively. Caution is therefore required when extrapolating from short‐term measurements to predict long‐term responses to global climate change. Our findings are of global significance as increases in atmospheric concentrations of CO₂ may, depending on sward composition and fertilizer management, increase greenhouse gas emissions of N₂O, thereby exacerbating the forcing effect of elevated CO₂ on global climate. Our results suggest that when applying high rates of N fertilizer to grassland systems, Trifolium repens swards, or a greater component of Trifolium in mixed swards, may minimize the negative effect of continued increasing atmospheric CO₂ concentrations on global warming.     

Physiological girdling of pine trees via phloem chilling: proof of concept

Quantifying below-ground carbon (C) allocation is particularly difficult as methods usually disturb the root-mycorrhizal-soil continuum. We reduced C allocation below ground of loblolly pine trees by: (1) physically girdling trees and (2) physiologically girdling pine trees by chilling the phloem. Chilling reduced cambium temperatures by approximately 18 degrees C. Both methods rapidly reduced soil CO2 efflux, and after approximately 10 days decreased net photosynthesis (P(n)), the latter indicating feedback inhibition. Chilling decreased soil-soluble C, indicating that decreased soil CO2 efflux may have been mediated by a decrease in root C exudation that was rapidly respired by microbes. These effects were only observed in late summer/early autumn when above-ground growth was minimal, and not in the spring when above-ground growth was rapid. All of the effects were rapidly reversed when chilling was ceased. In fertilized plots, both chilling and physical girdling methods reduced soil CO2 efflux by approximately 8%. Physical girdling reduced soil CO2 efflux by 26% in non-fertilized plots. This work demonstrates that phloem chilling provides a non-destructive alternative to reducing the movement of recent photosynthate below the point of chilling to estimate C allocation below ground on large trees.     

Below-ground respiratory responses of sugar maple and red maple saplings to atmospheric CO2 enrichment and elevated air temperature

The research described in this paper represents a part of a much broader research project with the general objective of describing the effects of elevated [CO2] and temperature on tree growth, physiological processes, and ecosystem-level processes. The specific objective of this research was to examine the below-ground respiratory responses of sugar maple (Acer saccharum Marsh.) and red maple (Acer rubrum L.) seedlings to elevated atmospheric [CO2] and temperature. Red maple and sugar maple seedlings were planted in the ground in each of 12 open-top chambers and exposed from 1994 through 1997 to ambient air or air enriched with 30 Pa CO2,< in combination with ambient or elevated (+4 °C) air temperatures. Carbon dioxide efflux was measured around the base of the seedlings and from root-exclusion zones at intervals during 1995 and 1996 and early 1997. The CO2 efflux rates averaged 0.4 μmol CO2 m-2 s-1 in the root-exclusion zones and 0.75 μmol CO2 m-2 s-1 around the base of the seedlings. Mineral soil respiration in root-exclusion zones averaged 12% higher in the high temperature treatments than at ambient temperature, but was not affected by CO2 treatments. The fraction of total efflux attributable to root + rhizosphere respiration ranged from 14 to 61% in measurements made around red maple plants, and from 35 to 62% around sugar maple plants. Root respiration rates ranged from 0 to 0.94 μmol CO2 s-1 m-2 of soil surface in red maple and from 0 to 1.02 in sugar maple. In both 1995 and 1996 root respiration rates of red maple were highest in high-CO2 treatments and lowest in high temperature treatments. Specific red maple root respiration rates of excised roots from near the soil surface in 1996 were also highest under CO2 enrichment and lowest in high temperature treatments. In sugar maple the highest rates of CO2 efflux were from around the base of plants exposed to both high temperature and high-CO2, even though specific respiration rates were< lowest for this species under the high temperature and CO2 enrichment regime. In both species, patterns of response to treatments were similar in root respiration and root mass, indicating that the root respiration responses were due in part to differences in root mass. The results underscore the need for separating the processes occurring in the roots from those in the forest floor and mineral soil in order to increase our understanding of the effects of global climate change on carbon sequestration and cycling in the below-ground systems of forests.     

Elevated [CO2], temperature increase and N supply effects on the turnover of below-ground carbon in a temperate grassland ecosystem

The effects of elevated [CO₂] (700 μl l^-1 CO₂) and temperature increase (+3 °C) on carbon turnover in grassland soils were studied during 2.5 years at two N fertiliser supplies (160 and 530 kg N ha^-1 y^-1) in an experiment with well-established ryegrass swards (Lolium perenne) supplied with the same amounts of irrigation water. During the growing season, swards from the control climate (350 μl l^-1 [CO₂] at outdoor air temperature) were pulse labelled by the addition of ¹³CO₂. The elevated [CO₂] treatments were continuously labelled by the addition of fossil-fuel derived CO₂ (^{13 C} of -40 to -50 ‰). Prior to the start of the experimental treatments, the carbon accumulated in the plant parts and in the soil macro-organic matter (‘old’ C) was at −32‰. During the experiment, the carbon fixed in the plant material (‘new’ C) was at −14 and −54‰ in the ambient and elevated [CO₂] treatments, respectively. During the experiment, the ¹³C isotopic mass balance method was used to calculate, for the top soil (0–15 cm), the carbon turnover in the stubble and roots and in the soil macro-organic matter above 200 μ (MOM). Elevated [CO₂] stimulated the turnover of organic carbon in the roots and stubble and in the MOM at N+, but not at N−. At the high N supply, the mean replacement time of ‘old’ C by ‘new’ C declined in elevated, compared to ambient [CO₂], from 18 to 7 months for the roots and stubble and from 25 to 17 months for the MOM. This resulted from increased rates of ‘new’ C accumulation and of ‘old’ C decay. By contrast, at the low N supply, despite an increase in the rate of accumulation of ‘new’ C, the soil C pools did not turnover faster in elevated [CO₂], as the rate of ‘old’ C decomposition was reduced. A 3 °C temperature increase in elevated [CO₂] decreased the input of fresh C to the roots and stubble and enhanced significantly the exponential rate for the ‘old’ C decomposition in the roots and stubble. An increased fertiliser N supply reduced the carbon turnover in the roots and stubble and in the MOM, in ambient but not in elevated [CO₂]. The respective roles for carbon turnover in the coarse soil OM fractions, of the C:N ratio of the litter, of the inorganic N availability and of a possible priming effect between C-substrates are discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Carbon and nitrogen allocation in Lolium perenne in response to elevated atmospheric CO2 with emphasis on soil carbon dynamics

The effect of elevated CO2 on the carbon and nitrogen distribution within perennial ryegrass (L. perenne L.) and its influence on belowground processes were investigated. Plants were homogeneously 14C-labelled in two ESPAS growth chambers in a continuous 14C-CO2 atmosphere of 350 and 700 L L-1 CO2 and at two soil nitrogen regimes, in order to follow the carbon flow through all plant and soil compartments.After 79 days, elevated CO2 increased the total carbon uptake by 41 and 21% at low (LN) and high nitrogen (HN) fertilisation, respectively. Shoot growth remained unaffected, whereas CO2 enrichment stimulated root growth by 46% and the root/soil respiration by 111%, irrespective of the nitrogen concentration. The total 14C-soil content increased by 101 and 28% at LN and HN, respectively. The decomposition of the native soil organic matter was not affected either by CO2 or by the nitrogen treatment.Elevated CO2 did not change the total nitrogen uptake of the plant either at LN or at HN. Both at LN and HN elevated CO2 significantly increased the total amount of nitrogen taken up by the roots and decreased the absolute and relative amounts translocated to the shoots.The amount of soil nitrogen immobilised by micro-organisms and the size of the soil microbial biomass were not affected by elevated CO2, whereas both were significantly increased at the higher soil N content.Most striking was the 88% increase in net carbon input into the soil expressed as: 14C-roots plus total 14C-soil content minus the 12C-carbon released by decomposition of native soil organic matter. The net carbon input into the soil at ambient CO2 corresponded with 841 and 1662 kg ha-1 at LN and HN, respectively. Elevated CO2 increased these amounts with an extra carbon input of 950 and 1056 kg ha-1. Combined with a reduced decomposition rate of plant material grown at elevated CO2 this will probably lead to carbon storage in grassland soils resulting in a negative feed back on the increasing CO2 concentration of the atmosphere.     

红豆草与土壤氮含量对大气二氧化碳浓度升高的响应

在封闭的植物培养箱中,通过盆栽实验,研究了红豆草和土壤氮含量对CO₂浓度增加的响应.结果表明,与正常CO₂浓度(355～370 μmol·mol^-1)相比,CO₂浓度升高(700 μmol·mol^-1),植物生物量增加25.1%(P＜0.01),但植物体氮浓度降低25.3%(P＜0.001),植物全氮没有显著的变化.经3个月盆栽实验后,与原始土壤相比,两种CO₂浓度处理土壤全N、NO₃^--N和NH₄⁺-N都有所降低,而土壤微生物氮则显著增加,这可能与植物生长有关.不同CO₂浓度处理土壤NH₄⁺-N浓度基本一致,但在高CO₂浓度下,土壤NO₃^--N浓度显著降低,而微生物生物氮显著增加.对整个土壤-植物系统而言,盆栽实验后,整个系统全氮有少量增加,但变化不显著,特别是在高CO₂浓度条件下,土壤-植物系统全氮最大,这可能与培养材料红豆草为豆科植物,而且在高CO₂浓度下生物量增加,导致氮的固定量增加有关.     

长期封育对不同类型草地碳贮量及其固持速率的影响

基于4个长期封育草地,采用成对取样方法(封育-自由放牧草地)分析了长期封育和自由放牧草地地上生物量、地表凋落物、0-100 cm根系和土壤的碳氮贮量,探讨了长期封育草地的碳固持速率。实验结果表明:长期封育显著提高了草地碳氮贮量;经30a围封处理后,草地碳固持量为1401-2858 g C m^-2,平均2126 g C m^-2;草地碳固持速率为46.7-129.2 g C m^-2 a^-1,平均84.2 g C m^-2 a^-1。长期封育草地氮固持速率为2.8-14.7 g N m^-2 a^-1,平均7.3 g N m^-2 a^-1。封育草地碳和氮固持速率表现为:针茅草地＜羊草草地＜退化羊草草地＜补播黄花苜蓿+羊草草地。长期封育草地0-40 cm土壤碳固持速率相对较高,但下层土壤对草地碳固持的贡献也比较大,因此,未来的相关研究应给予下层土壤更大关注。内蒙古典型草地具有巨大的碳固持潜力,长期封育(或禁牧)是实现其碳固持效应最经济、最有效的途径之一。     

干扰对典型草原生态系统土壤净呼吸特征的影响

由于土地利用格局的改变和人类干扰活动的加剧,草地生态系统CO2排放与固定的平衡、碳循环特征以及碳储量越来越受到人们的重视。尤其是定量区分土壤净呼吸与土壤总呼吸量之间的比例关系,以及定量描述草地生态系统碳循环过程等方面的研究尚不够完善。以河北沽源的典型草原为研究对象,测定了火烧、灌溉、施肥、刈割干扰下的天然草地土壤净呼吸变化动态及其与主要控制因素之间的关系。结果表明:不同处理土壤净呼吸均表现出明显的季节性变化规律,变化趋势基本一致。火烧、灌溉和刈割处理分别比对照的土壤净呼吸通量降低了28.93%、16.25%和36.82%。土壤温度、土壤湿度与土壤净呼吸通量呈指数相关(P0.01)。对地上生物量、地下生物量、土壤有机碳含量和土壤全氮含量与土壤净呼吸之间进行逐步回归分析表明,土壤有机碳含量(SC)和土壤全氮含量(SN)是土壤净呼吸通量的主要影响因素。     

Soil nutrient heterogeneity interacts with elevated CO2 and nutrient availability to determine species and assemblage responses in a model grassland community

Interactive effects of atmospheric CO(2) concentration ([CO(2)]), soil nutrient availability and soil nutrient spatial distribution on the structure and function of plant assemblages remain largely unexplored. Here we conducted a microcosm experiment to evaluate these interactions using a grassland assemblage formed by Lolium perenne, Plantago lanceolata, Trifolium repens, Anthoxanthum odoratum and Holcus lanatus. Assemblages exhibited precise root foraging patterns, had higher total and below-ground biomass, and captured more nitrogen when nutrients were supplied heterogeneously. Root foraging responses were modified by nutrient availability, and the patterns of N capture by interactions between nutrient distribution, availability and [CO(2)]. Greater above-ground biomass was observed under elevated CO(2) only under homogeneous conditions of nutrient supply and at the highest availability level. CO(2) interacted with nutrient distribution and availability to determine foliar percentage N and below : above-ground biomass ratios, respectively. Interactions between nutrient distribution and CO(2) determined the relative contribution to above-ground biomass of four of the species. The responses of dominant and subordinate species to [CO(2)] were dependent on the availability and distribution of nutrients. Our results suggest that soil nutrient distribution has the potential to influence the response of plant species and assemblages to changes in [CO(2)] and nutrient availability.     

内蒙古放牧草地土壤碳固持速率和潜力

放牧是典型草地最重要的利用方式,弄清放牧对草地碳固速率的影响,将为我国内蒙古地区草地碳汇管理提供重要的科学依据。通过在平坦草地和斜坡草地设置相同的放牧梯度实验 (放牧强度0、1.5、3.0、4.5、6.0、7.5、9.0 羊/hm²),探讨了放牧和地形对草地土壤碳固持速率的影响。实验结果表明:轻度放牧草地表现为碳固持,重度放草地表现为碳流失;对放牧草地而言,存在碳源/碳汇的转化阈值(或放牧强度),且坡地阈值低于平地。为了实现草地碳增汇目的,平坦草地的放牧强度应低于 4.5羊/hm²(放牧期6-9月),斜坡草地应低于3 羊/hm²。地形因素(平地VS斜坡)使准确评估放牧草地土壤的碳固持速率变得更加复杂。总之,内蒙古地区放牧草地具有较大的碳固持潜力,通过控制放牧强度是实现其碳固持潜力的重要途径之一。     

Carbon‐13 input and turn‐over in a pasture soil exposed to long‐term elevated atmospheric CO2

The impact of elevated CO₂ and N‐fertilization on soil C‐cycling in Lolium perenne and Trifolium repens pastures were investigated under Free Air Carbon dioxide Enrichment (FACE) conditions. For six years, swards were exposed to ambient or elevated CO₂ (35 and 60 Pa pCO₂) and received a low and high rate of N fertilizer. The CO₂ added in the FACE plots was depleted in ¹³C compared to ambient (Δ? 40‰) thus the C inputs could be quantified. On average, 57% of the C associated with the sand fraction of the soil was ‘new’ C. Smaller proportions of the C associated with the silt (18%) and clay fractions (14%) were derived from FACE. Only a small fraction of the total C pool below 10 cm depth was sequestered during the FACE experiment. The annual net input of C in the FACE soil (0–10 cm) was estimated at 4.6 ± 2.2 and 6.3 ± 3.6 (95% confidence interval) Mg ha^{? 1} for T. repens and L. perenne, respectively. The maximum amount of labile C in the T. repens sward was estimated at 8.3 ± 1.6 Mg ha^{? 1} and 7.1 ± 1.0 Mg ha^{? 1} in the L. perenne sward. Mean residence time (MRT) for newly sequestered soil C was estimated at 1.8 years in the T. repens plots and 1.1 years for L. perenne. An average of 18% of total soil C in the 0–10 cm depth in the T. repens sward and 24% in the L. perenne sward was derived from FACE after 6 years exposure. The majority of the change in soil δ¹³C occurred in the first three years of the experiment. No treatment effects on total soil C were detected. The fraction of FACE‐derived C in the L. perenne sward was larger than in the T. repens sward. This suggests a priming effect in the L. perenne sward which led to increased losses of the old C. Although the rate of C cycling was affected by species and elevated CO₂, the soil in this intensively managed grassland ecosystem did not become a sink for additional new C.     

Total soil C and N sequestration in a grassland following 10 years of free air CO2 enrichment

Soil C sequestration may mitigate rising levels of atmospheric CO_2. However, it has yet to be determined whether net soil C sequestration occurs in N‐rich grasslands exposed to long‐term elevated CO₂. This study examined whether N‐fertilized grasslands exposed to elevated CO₂ sequestered additional C. For 10 years, Lolium perenne, Trifolium repens, and the mixture of L. perenne/T. repens grasslands were exposed to ambient and elevated CO₂ concentrations (35 and 60 Pa pCO₂). The applied CO₂ was depleted in δ¹³C and the grasslands received low (140 kg ha^?1) and high (560 kg ha^?1) rates of ¹⁵N‐labeled fertilizer. Annually collected soil samples from the top 10 cm of the grassland soils allowed us to follow the sequestration of new C in the surface soil layer. For the first time, we were able to collect dual‐labeled soil samples to a depth of 75 cm after 10 years of elevated CO₂ and determine the total amount of new soil C and N sequestered in the whole soil profile. Elevated CO₂, N‐fertilization rate, and species had no significant effect on total soil C. On average 9.4 Mg new C ha^?1 was sequestered, which corresponds to 26.5% of the total C. The mean residence time of the C present in the 0–10 cm soil depth was calculated at 4.6±1.5 and 3.1±1.1 years for L. perenne and T. repens soil, respectively. After 10 years, total soil N and C in the 0–75 cm soil depth was unaffected by CO₂ concentration, N‐fertilization rate and plant species. The total amount of ¹⁵N‐fertilizer sequestered in the 0–75 cm soil depth was also unaffected by CO₂ concentration, but significantly more ¹⁵N was sequestered in the L. perenne compared with the T. repens swards: 620 vs. 452 kg ha^?1 at the high rate and 234 vs. 133 kg ha^?1 at the low rate of N fertilization. Intermediate values of ¹⁵N recovery were found in the mixture. The fertilizer derived N amounted to 2.8% of total N for the low rate and increased to 8.6% for the high rate of N application. On average, 13.9% of the applied ¹⁵N‐fertilizer was recovered in the 0–75 cm soil depth in soil organic matter in the L. perenne sward, whereas 8.8% was recovered under the T. repens swards, indicating that the N₂‐fixing T. repens system was less effective in sequestering applied N than the non‐N₂‐fixing L. perenne system. Prolonged elevated CO₂ did not lead to an increase in whole soil profile C and N in these fertilized pastures. The potential use of fertilized and regular cut pastures as a net soil C sink under long‐term elevated CO₂ appears to be limited and will likely not significantly contribute to the mitigation of anthropogenic C emissions.     

CO2浓度升高和施氮条件下小麦根际呼吸对土壤呼吸的贡献

依托FACE技术平台,采用稳定13C同位素技术,通过将小麦(C3作物)种植于长期单作玉米(C4作物)的土壤上,研究了大气CO2浓度升高和不同氮肥水平对土壤排放CO2的δ13C值及根际呼吸的影响.结果表明:种植小麦后土壤排放CO2的δ13C值随作物生长逐渐降低,CO2浓度升高200 μmol·mol-1显著降低了孕穗、抽穗期(施氮量为250 kg·hm-2,HN)与拔节、孕穗期(施氮量为150 kg·hm-2,LN)土壤排放CO2的δ13C值,显著提高了孕穗、抽穗期的根际呼吸比例.拔节至成熟期,根际呼吸占土壤呼吸的比例在高CO2浓度下为24％～48％ (HN)和21％ ～48％ (LN),在正常CO2浓度下为20％ ～36％ (HN)和19％～32％(LN).不同CO2浓度下土壤排放CO2的δ13C值和根际呼吸对氮肥增加的响应不同,CO2浓度与氮肥用量在拔节期对根际呼吸的交互效应显著.     

Elevated CO2 increases nitrogen rhizodeposition and microbial immobilization of root-derived nitrogen

With this study, we aimed to determine how elevated CO(2) affects rhizodeposition and the cycling of rhizodeposited nitrogen (N) in the soil under C(3) and C(4) plants. In addition, we examined how cultivated genotypes of wheat (Triticum turgidum) and maize (Zea mays) responded to elevated CO(2) in comparison with their wild relatives. By constructing an N-transfer experiment we could directly assess cycling of the rhizodeposited N and trace the fate of rhizodeposited N in the soil and in receiver plants. Biomass production, rhizodeposition and cycling of root-borne N in maize genotypes were not affected by elevated CO(2). Elevated CO(2) stimulated above- and below-ground biomass production of the wheat genotypes on average by 38%, and increased rhizodeposition and immobilization of root-derived N on average by 30%. Concurrently, elevated CO(2) reduced mineral (15)N and re-uptake of the root-derived N by 50% in wheat. This study shows that elevated CO(2) may enhance N limitation by increasing N rhizodeposition and subsequent immobilization of the root-derived N.