氮沉降对森林生态系统碳吸存的影响

工业化带来的大气氮沉降增加是影响森林生态系统碳吸存的重要因素。将森林碳库分为地上和地下两部分,从3个方面综述了国内外氮沉降对森林生态系统碳吸存影响的研究现状。(1)地上部分:氮限制的温带森林,氮沉降增加了地上部分碳吸存。氮丰富的热带森林,氮沉降对地上部分碳吸存没有影响。过量的氮输入会造成森林死亡率的上升,从而降低地上部分碳吸存。(2)地下部分:相比地上部分研究得少,表现为增加、降低和没有影响3种效果。(3)目前的结论趋向于认为氮沉降促进森林生态系统碳吸存,然而氮沉降所带来的森林生态系统碳吸存能力到底有多大依然无法确定,这也将成为未来氮碳循环研究的重点问题。分析了氮沉降影响森林生态系统碳吸存的机理,介绍了氮沉降对森林生态系统碳吸存影响的4种研究方法。探讨了该领域研究的不足及未来的研究方向。     

氮沉降对森林生物多样性的影响

从3个方面论述了氮沉降对森林生物多样性影响：（1）森林植物多样性,包括乔木层植物、林下层植物和隐花植物;（2）土壤微生物多样性,主要是细菌和真菌;（3）森林动物多样性：主要包括地下土壤动物和地上草食动物。综合来看,氮沉降改变了物种组成,过量氮沉降降低了生物多样性。同时,也对氮沉降影响生物多样性的机理进行了分析。最后,还探讨了当前在氮沉降对森林生物多样性影响的研究方面存在的问题以及今后研究的方向。     

大气氮沉降与森林生态系统的氮动态

由于人类活动的影响,若干年代以来大气氮沉降明显增加。在森林地区,大气氮沉降的空间变异性由林分的位置、结构和组成树种所决定。除降雨之外,干沉降和隐藏降水也是大气氮沉降的重要形式。     

陆地生态系统氮沉降增加的生态效应

 人类活动在全球范围内极大地改变着氮素从大气向陆地生态系统输入的方式和速率,人为固定的氮素正在不断积累,并对生态系统的结构和功 能产生显著影响。该文从以下几个方面综述了大气氮沉降增加对陆地生态系统的影响：1)氮输入增加可能影响植物生产力和生态系统碳蓄积能 力,生态系统响应的方向和程度取决于系统的初始氮状况(氮限制或氮饱和)以及当地的植被和土壤特征;2)持续氮输入有可能改变土壤氮循环 过程,降低土壤固持氮的能力,甚至导致土壤酸化、盐基离子损耗,进而影响到土壤有机碳的分解;3)高的氮沉降速率和持续氮输入都可能加 速含氮痕量气体的释放,但其影响程度受生态系统初始状态的影响(例如磷限制和氮限制);4)氮沉降增加会影响生态系统的物种丰富度、植物 群落结构和动态,促进森林扩张,改变菌根真菌的物种多样性;5)持续氮输入带来的植物群落结构和植物生理特征的变化可能影响昆虫取食特 性,进而通过食物链改变生态系统的营养结构;6) 氮沉降增加对生态系统的影响并不是孤立存在的,它与CO₂浓度升高和O₃浓度变化有协同作 用,但难以从其协同效应中区分出各自的影响。最后,该文总结了我国的氮沉降研究现状,并对今后的研究前景提出了展望。     

模拟大气沉降氮在陆地生态系统中的留存及其影响因素

大气氮沉降对陆地生态系统的影响取决于氮在生态系统中的留存量。探明生态系统的氮留存作用对预测全球变化背景下生态系统氮循环具有重要意义,但植物、土壤和生态系统氮留存的影响因素仍不清楚。通过文献搜集对58个野外¹⁵N同位素示踪研究的305组观测结果进行综合分析。结果表明:生态系统氮回收率(¹⁵N_rec)为(56.3±1.39)%,且土壤¹⁵N_rec (40.1±1.17)%显著高于植物¹⁵N_rec (16.2±0.89)% (P < 0.001)。年均温度、年降水量、土壤有机碳、总氮和碳氮比的增加显著增加植物、土壤和生态系统¹⁵N_rec,但土壤pH、施氮量和施氮持续时间的增加却显著降低植物和生态系统¹⁵N_rec (P < 0.05)。上述因素解释了生态系统、植物和土壤¹⁵N_rec变化的65.0%,61.0%和64.0%。年降水量是影响生态系统、植物和土壤¹⁵N_rec的最主要因素(P < 0.001),年降水量和年均温度构成的气候因素显著影响生态系统、植物和土壤¹⁵N_rec。研究结果对阐明陆地生态系统氮循环过程,以及维持生态系统的稳定性具有重要意义。     

长白山森林生态系统大气氮素湿沉降通量和组成的季节变化特征

2009—2010年期间,利用雨量计收集法在长白山森林生态系统定位站开展定位观测,分析降水中氮素浓度,研究了该区域大气氮素湿沉降通量和组成的季节变化特征。结果表明,各形态氮素月均浓度之间差别较大,具有明显的季节性;其降水中浓度主要受降水量和降水频次的影响。全年氮素湿沉降中TN、TIN和TON的沉降量分别为27.64 kg N hm-2a-1、11.05 kg N hm-2a-1和16.59 kg N hm-2a-1,TON为沉降主体,占60.02%;其大气氮沉降量主要由降水量和降水中氮素浓度共同决定。该地区氮湿沉降量已处于我国中等水平,考虑到氮素的干湿沉降比例,本区域的年氮沉降量已接近或超过本区域的营养氮沉降临界负荷,存在一定的环境风险。该地区生长季(5—10月)的氮沉降量(16.59 kg N hm-2a-1)占全年氮沉降量的比例达到73.20%。生长季的氮沉降对于促进植物生长直接生态意义重大,而非生长季的氮沉降对于大量补充次年植物生长初期所需养分的间接生态意义明显。     

森林生态系统碳循环对全球氮沉降的响应

森林土壤和植被储存着全球陆地生态系统大约46%的碳,在全球碳平衡中起着非常重要的作用。过去几十年来,森林生态系统的碳循环和碳吸存受到了全球氮沉降的深刻影响,因为氮沉降改变了陆地生态系统的生产力和生物量积累。以欧洲和北美温带森林区域开展的研究为基础,综述了氮沉降对植物光合作用、土壤呼吸、土壤DOM及林木生长的影响特征和机理,探讨了森林生态系统碳动态对氮沉降响应的不确定性因素。热带森林C、N循环与大部分温带森林不同,人为输入的氮对热带生态系统过程的影响也可能不同,因此指出了在热带地区开展碳氮循环耦合研究的必要性和紧迫性。     

氮沉降对森林植物的影响

综述了氮沉降对森林植物的影响。氮沉降对森林植物的影响主要表现在以下6个方面：(1)在一定量范围内的氮沉降有利于植物的光合作用,但过量后则会引起植物的光合速率下降;(2)当植物生长受氮限制时,在一定程度上的氮沉降增加植物生产力,但当氮过量后,氮沉降则使植物的生产力下降;(3)过量的氮沉降导致植物体各种营养元素含量的比例失衡;(4)氮沉降会改变植物的形态结构,集中表现为根／冠比减小;(5)氮沉降会增加植物对天然胁迫如干旱、病虫害和风的敏感性,减少其抵御能力;(6)氮沉降会改变植物组成和降低森林植物的多样性。     

大气N沉降的不断增加对森林生态系统的影响

若干年代以来,大气Ｎ沉降不断增加．在一些地区,大气Ｎ沉降超过了森林生态系统的Ｎ需求．Ｎ沉降的增加对植物生长的刺激作用和对菌根的危害、过剩的ＮＨ４＋在体内对其它阳离子的交换取代和在土壤中对其它阳离子在根的养分吸收方面的竞争抑制,都可造成植物体内其它养分缺乏,导致森林营养失调．Ｎ沉降的增加将提高硝化作用,加速ＮＯ３－和盐基阳离子的淋失,引起土壤酸化和Ａｌ、Ｍｎ活化．植物体内的高Ｎ水平将增加森林对寒冷、霜冻、真菌病害及可能的虫害等胁迫的敏感性．Ｎ沉降长期而持续的增加可通过干扰演替动力学,促使植物群落发生变化．     

氮沉降对森林生态系统土壤碳库的影响

森林土壤碳库是陆地生态系统碳库的重要组成部分,对维持全球碳平衡具有重要意义。不断加剧的全球氮沉降有可能改变森林生态系统中碳元素的地球化学循环过程,从而引起森林土壤碳储量的变化。本文从森林土壤碳收支的角度,将氮沉降对森林生态系统土壤碳库影响的复杂过程划分为凋落物分解、细根周转、土壤呼吸和土壤可溶性有机碳淋失4个相对独立的过程。综合国内外研究现状,对其进行了简要评述,指出了目前研究的不足,并探讨了这一研究领域的发展方向。     

Distribution of nitrogen-15 tracers applied to the canopy of a mature spruce-hemlock stand,Howland, Maine,USA

In N-limited ecosystems, fertilization by N deposition may enhance plant growth and thus impact C sequestration. In many N deposition–C sequestration experiments, N is added directly to the soil, bypassing canopy processes and potentially favoring N immobilization by the soil. To understand the impact of enhanced N deposition on a low fertility unmanaged forest and better emulate natural N deposition processes, we added 18 kg N ha⁻¹ year⁻¹ as dissolved NH₄NO₃ directly to the canopy of 21 ha of spruce-hemlock forest. In two 0.3-ha subplots, the added N was isotopically labeled as ¹⁵NH₄ ⁺ or ¹⁵NO₃ ⁻ (1% final enrichment). Among ecosystem pools, we recovered 38 and 67% of the ¹⁵N added as ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻, respectively. Of ¹⁵N recoverable in plant biomass, only 3–6% was recovered in live foliage and bole wood. Tree twigs, branches, and bark constituted the most important plant sinks for both NO₃ ⁻ and NH₄ ⁺, together accounting for 25–50% of ¹⁵N recovery for these ions, respectively. Forest floor and soil ¹⁵N retention was small compared to previous studies; the litter layer and well-humified O horizon were important sinks for NH₄ ⁺ (9%) and NO₃ ⁻ (7%). Retention by canopy elements (surfaces of branches and boles) provided a substantial sink for N that may have been through physico-chemical processes rather than by N assimilation as indicated by poor recoveries in wood tissues. Canopy retention of precipitation-borne N added in this particular manner may thus not become plant-available N for several years. Despite a large canopy N retention potential in this forest, C sequestration into new wood growth as a result of the N addition was only ~16 g C m⁻² year⁻¹ or about 10% above the current net annual C sequestration for this site.     

Elevated [CO2] and increased N supply reduce leaf disease and related photosynthetic impacts on Solidago rigida

To evaluate whether leaf spot disease and related effects on photosynthesis are influenced by increased nitrogen (N) input and elevated atmospheric CO₂ concentration ([CO₂]), we examined disease incidence and photosynthetic rate of Solidago rigida grown in monoculture under ambient or elevated (560 μmol mol⁻¹) [CO₂] and ambient or elevated (+4 g N m⁻² year⁻¹) N conditions in a field experiment in Minnesota, USA. Disease incidence was lower in plots with either elevated [CO₂] or enriched N (−57 and −37%, respectively) than in plots with ambient conditions. Elevated [CO₂] had no significant effect on total plant biomass, or on photosynthetic rate, but reduced tissue%N by 13%. In contrast, N fertilization increased both biomass and total plant N by 70%, and as a consequence tissue%N was unaffected and photosynthetic rate was lower on N fertilized plants than on unfertilized plants. Regardless of treatment, photosynthetic rate was reduced on leaves with disease symptoms. On average across all treatments, asymptomatic leaf tissue on diseased leaves had 53% lower photosynthetic rate than non-diseased leaves, indicating that the negative effect from the disease extended beyond the visual lesion area. Our results show that, in this instance, indirect effects from elevated [CO₂], i.e., lower disease incidence, had a stronger effect on realized photosynthetic rate than the direct effect of higher [CO₂].     

Ecological Footprint: Refining the carbon Footprint calculation

Within the Ecological Footprint methodology, the carbon Footprint component is defined as the regenerative forest capacity required to sequester the anthropogenic carbon dioxide emissions that is not absorbed by oceans. A key parameter of the carbon Footprint is the Average Forest Carbon Sequestration (AFCS), which is calculated from the net carbon sequestration capacity of forests ecosystems.The aim of this paper is to increase the clarity and transparency of the Ecological Footprint by reviewing the rationale and methodology behind the carbon Footprint component, and updating a key factor in its calculation, the AFCS. Multiple calculation options have been set to capture different rates of carbon sequestration depending on the degree of human management of three types of forest considered (primary forests, other naturally regenerated forests and planted forests). Carbon emissions related to forest wildfires and soil as well as harvested wood product have been included for the first time in this update of the AFCS calculation. Overall, a AFCS value range of 0.73 ± 0.37 t C ha⁻¹ yr⁻¹ has been identified. The resulting carbon Footprint and Ecological Footprint values have then been evaluated based on this value range. Results confirm that human demand for ecosystem services is beyond the biosphere's natural capacity to provide them.     

Elevated CO2 increases belowground respiration in California grasslands

This study was designed to identify potential effects of elevated CO₂ on belowground respiration (the sum of root and heterotrophic respiration) in field and microcosm ecosystems and on the annual carbon budget. We made three sets of respiration measurements in two CO₂ treatments, i.e., (1) monthly in the sandstone grassland and in microcosms from November 1993 to June 1994; (2) at the annual peak of live biomass (March and April) in the serpentine and sandstone grasslands in 1993 and 1994; and (3) at peak biomass in the microcosms with monocultures of seven species in 1993. To help understand ecosystem carbon cycling, we also made supplementary measurements of belowground respiration monthly in sandstone and serpentine grasslands located within 500 m of the CO₂ experiment site. The seasonal average respiration rate in the sandstone grassland was 2.12 mol m^-2 s^-1 in elevated CO₂, which was 42% higher than the 1.49 mol m^-2 s^-1 measured in ambient CO₂ (P=0.007). Studies of seven individual species in the microcosms indicated that respiration was positively correlated with plant biomass and increased, on average, by 70% with CO₂. Monthly measurements revealed a strong seasonality in belowground respiration, being low (0–0.5 mol CO₂ m^-2 s^-1 in the two grasslands adjacent to the CO₂ site) in the summer dry season and high (2–4 mol CO₂ m^-2 s^-1 in the sandstone grassland and 2–7 mol CO₂ m^-2 s^-1 in the microcosms) during the growing season from the onset of fall rains in November to early spring in April and May. Estimated annual carbon effluxes from the soil were 323 and 440 g C m^-2 year^-1 for the sandstone grasslands in ambient and elevated CO₂. That CO₂-stimulated increase in annual soil carbon efflux is more than twice as big as the increase in aboveground net primary productivity (NPP_a) and approximately 60% of NPP_a in this grassland in the current CO₂ environment. The results of this study suggest that below-ground respiration can dissipate most of the increase in photosynthesis stimulated by elevated CO₂.CIWDPB Publication # 1271     

Non-structural carbohydrate pools in a tropical forest

The pool size of mobile, i.e. non-structural carbohydrates (NSC) in trees reflects the balance between net photosynthetic carbon uptake (source) and irreversible investments in structures or loss of carbon (sink). The seasonal variation of NSC concentration should reflect the sink/source relationship, provided all tissues from root to crown tops are considered. Using the Smithsonian canopy crane in Panama we studied NSC concentrations in a semi-deciduous tropical forest over 22 months. In the 9 most intensively studied species (out of the 17 investigated), we found higher NSC concentrations (starch, glucose, fructose, sucrose) across all species and organs in the dry season than in the wet season (NSC 7.2% vs 5.8% of dry matter in leaves, 8.8/6.0 in branches, 9.7/8.5 in stems, 8.3/6.4 in coarse and 3.9/2.2 in fine roots). Since this increase was due to starch only, we attribute this to drought-constrained growth (photosynthesis less affected by drought than sink activity). Species-specific phenological rhythms (leafing or fruiting) did not overturn these seasonal trends. Most of the stem volume (diameter at breast height around 40 cm) stores NSC. We present the first whole forest estimate of NSC pool size, assuming a 200 t ha^–1 forest biomass: 8% of this i.e. ca. 16 t ha^–1 is NSC, with ca. 13 t ha^–1 in stems and branches, ca. 0.5 and 2.8 t ha^–1 in leaves and roots. Starch alone (ca. 10.5 t ha^–1) accounts for far more C than would be needed to replace the total leaf canopy without additional photosynthesis. NSC never passed through a period of significant depletion. Leaf flushing did not draw heavily upon NSC pools. Overall, the data imply a high carbon supply status of this forest and that growth during the dry season is not carbon limited. Rather, water shortage seems to limit carbon investment (new tissue formation) directly, leaving little leeway for a direct CO₂ fertilization effects.     

Early stage litter decomposition rates for Swiss forests

The decomposition of belowground and aboveground tree litter was studied on five forest sites across Switzerland, ranging from 480 to 1500 m in altitude, and including calcareous and acidic soils. In addition to decomposition of local litter types (Picea abies, Fagus sylvatica, Castanea sativa), the decomposition of a standard beech litter was studied on all sites. After 2 years of decomposition, mass loss ranged from 18 to 71% across the different sites and litter types. The lowest decomposition rates were observed for beech roots, while mass loss was greatest for both spruce needles and spruce roots at the low-altitude site. Mass loss during the first winter correlated best with the content of water-soluble substances. After 1 year of incubation, mass loss of the standard litter varied less than did mass loss of local litter, but variance increased during the second year for aboveground litter. These observations indicate a smaller climatic influence on litter breakdown at the beginning of the decomposition process. Litter mass loss could be described using an exponential model with a decay constant depending on either lignin/N ratio or Mn content of the litter and annual soil temperature and throughfall precipitation as climatic variables. Modelling the observed mass loss indicated a strong influence of litter quality in the first 2 years of decomposition, confirming the field data from the standard litter experiment. The experiment will continue for some years and is expected to yield additional data on long-term decomposition.     

开放系统中农作物对空气CO2浓度增加的响应

FACE试验（free-air CO2 enrichment)开展的10多年中,供试农作物主要有：C3禾本科作物小麦（Triticum aestivum L.）、多年生黑麦草（Lolium perenne)和水稻（Oryza sativaL.),C4禾本科类高梁（Sorghum bicolor(L.)Moench),C3豆科植物白三叶草（Trifolium repens ),C3非禾本科块茎状作物马铃薯（Solanum tuberosum L.）,以及多年生C3类木作物棉花（Gossypium hirsutum L.)和葡萄（Vitisvinifera l.)。本文系统整理和分析了以下各项参数的结果;光合作用、气孔导度、冠层温度、水分利用、水势、叶面积指数、根茎生物量累积、作物产量、辐射利用率,比叶面积、N含量、N收益、碳水化合物含量、物候变化、土壤微生物、土壤呼吸、痕量气体交换以及土壤碳固定,CO2浓度升高对农作物的影响作用主要表现在以下方面：（1）促进了植物光合作用,增加了其生物量累积;（2）显著提高C3作物产量,但对C4作物产量的影响很小;（3）降低了C3和C4作物气孔导度,非常显著地提高了所有作物的水分利用率;（4）对植物生长的促进作用在水分不足与水分充中时二者相当或前者大于后者;（6）对根系生长的促进作用要大于地上部分;（7）对多年生植物气孔导度的影响较小,但对其生长的促进作用仍很高;（8）降低了植物体内N含量,但作物体内碳水化合物及某些其他含碳化合物含量增加,且叶部含量要明显高于植物其他器官;（9）对大多数作物的物候略有加速;（10）对某些土壤微生物具显著影响,而对有些则无,但都增加了微生物活性;（11）综合多年、多地点的试验结果表明土壤对大气CO2的固定增加,但单独一个试验无法观测到SOC的显著性变化,对FACE和前期的熏气室试验结果都进行了尽可能的对比研究,除了二例以外,发现在大多数情况下二者的结果基本一致,其中,FACE使气孔导度降低的1.5倍,明显高于前期熏气室试验的结果;其二,相对于熏气室,FACE条件下CO2倍增对根的相对促进作用要高于地上部分,因此,我们对基于这二者的结论的准确性和可靠性是充满信心的,不过,更接近自然环境和具更大小区面积的FACE试验仍是必需的,它可以为我们提供在CO2升高条件下更具代表性的田间试验条件,从而为我们提供更多、更有益的多学科交叉的试验数据和研究结果。     

Nitrogen supply effects on productivity and potential leaf litter decay of Carex species from peatlands differing in nutrient limitation

We investigated the effect of increased N-supply on productivity and potential litter decay rates of Carex species, which are the dominant vascular plant species in peatlands in the Netherlands. We hypothesized that: (1) under conditions of N-limited plant growth, increased N-supply will lead to increased productivity but will not affect C:N ratios of plant litter and potential decay rates of that litter; and (2) under conditions of P-limited plant growth, increased N-supply will not affect productivity but it will lead to lower C:N ratios in plant litter and thereby to a higher potential decay rate of that litter. These hypotheses were tested by fertilization experiments (addition of 10 g N m^-2 year^-1) in peatlands in which plant growth was N-limited and P-limited, respectively. We investigated the effects of fertilization on net C-fixation by plant biomass, N uptake, leaf litter chemistry and potential leaf litter decay. In a P-limited peatland, dominated by Carex lasiocarpa, there was no significant increase of net C-fixation by plant biomass upon enhanced N-supply, although N-uptake had increased significantly compared with the unfertilized control. Due to the N-fertilization the C:N ratio in the plant biomass decreased significantly. Similarly, the C:N ratio of leaf litter produced at the end of the experiment showed a significant decrease upon enhanced N-supply. The potential decay rate of that litter, measured as CO₂-evolution from the litter under aerobic conditions, was significantly increase upon enhanced N-supply. In a N-limited peatland, dominated by C. acutiformis, the net C-fixation by plant biomass increased with increasing N-supply, whereas the increase in N-uptake was not significant. The C:N ratio of both living plant material and of dead leaves did not change in response to N-fertilization. The potential decay rate of the leaf litter was not affected by N-supply. The results agree with our hypotheses. This implies that atmospheric N-deposition may affect the CO₂-sink function of peatlands, but the effect is dependent on the nature of nutrient limitation. In peatlands where plant growth is N-limited, increased N-supply leads to an increase in the net accumulation of C. Under conditions of P-limited plant growth, however, the net C-accumulation will decrease, because productivity is not further increased, whereas the amount of C lost through decomposition of dead organic matter is increased. As plant growth in most terrestrial ecosystems is N-limited, increased N-supply will in most peatlands lead to an increase of net C-accumulation.