森林土壤氮素转换及其对氮沉降的响应

近几十年人类活动向大气中排放的含氮化合物激增 ,并引起大气氮沉降也成比例增加。目前 ,氮沉降的增加使一些森林生态系统结构和功能发生改变 ,甚至衰退。近 2 0 a欧洲和北美有关氮沉降及其对森林生态系统的影响方面的研究较多 ,而我国少有涉及。森林土壤氮素转换是森林生态系统氮素循环的一个重要的组成部分 ,而矿化、硝化和反硝化作用是其核心过程 ,氮沉降作为驱动因子势必改变森林土壤氮素转换速度、方向和通量。根据国外近 2 0 a有关研究 ,首先介绍了森林土壤氮素转换过程和强度 ,论述森林土壤氮素在生态系统氮素循环中的作用 ,然后在此基础上 ,介绍了氮沉降对森林土壤氮素循环的研究途径 ,探讨了氮沉降对森林土壤氮素矿化、硝化和反硝化作用的影响及其机理     

Major changes in forest carbon and nitrogen cycling caused by declining sulphur deposition

Sulphur (S) and nitrogen (N) deposition are important drivers of the terrestrial carbon (C) and N cycling. We analyzed changes in C and N pools in soil and tree biomass at a highly acidified spruce site in the Czech Republic during a 15 year period. Total S deposition decreased from 5 to 1.1 g m^?2 yr^?1 between 1995 and 2009, whereas bulk N deposition did not change. Over the same period, C and N pools in the Oa horizon declined by 116 g C and 4.2 g N m^?2 yr^?1, a total decrease of 47% and 42%, respectively. This loss of C and N probably originated from organic matter (OM) that had accumulated during the period of high acid deposition when litter decomposition was suppressed. The loss of OM from the Oa horizon coincided with a substantial leaching (1.3 g N m^?2 yr^?1 at 90 cm) in the 1990s to almost no leaching (<0.02 g N m^?2 yr^?1) since 2006. Forest floor net N mineralization also decreased. This had consequences for spruce needle N concentration (from 17.1 to 11.4 mg kg^?1 in current needles), an increase in litterfall C/N ratio (from 51 to 63), and a significant increase in the Oi + Oe horizon C/N ratio (from 23.4 to 27.3) between 1994 and 2009/2010. Higher forest growth and lower canopy defoliation was observed in the 2000s compared to the 1990s. Our results demonstrate that reducing S deposition has had a profound impact on forest organic matter cycling, leading to a reversal of historic ecosystem N enrichment, cessation of nitrate leaching, and a major loss of accumulated organic soil C and N stocks. These results have major implications for our understanding of the controls on both N saturation and C sequestration in forests, and other ecosystems, subjected to current or historic S deposition.     

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。     

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。     

森林土壤微生物对干旱和氮沉降的响应

干旱和氮沉降深刻影响着人类世森林生态系统的生命活动与物质循环,进而影响全球碳平衡、并反馈作用于气候变化。土壤微生物驱动元素的生物地球化学循环和关键土壤生态过程,在气候变化生物学研究方面具有核心地位和全球重要性。本文综述了干旱和氮沉降对森林土壤细菌和菌根真菌的影响。提出未来应加强全球变化多因子交互作用对土壤微生物多样性、活性与生态功能的研究;建立野外长期定位站,强化亚热带森林生态系统与全球变化研究;注重土壤生物之间互作及网络研究;利用微生物大数据建立相关的机理模型等。从认识微生物多样性和群落组成对全球变化的响应与适应,逐步发展为调控利用微生物群落服务于森林的优化管理、生态资源的合理保护与可持续利用,为充分发挥微生物减缓全球气候变化的作用提供理论基础。     

Regional scale gradients of climate and nitrogen deposition drive variation in ectomycorrhizal fungal communities associated with native Scots pine

Ectomycorrhizal fungi commonly associate with the roots of forest trees where they enhance nutrient and water uptake, promote seedling establishment and have an important role in forest nutrient cycling. Predicting the response of ectomycorrhizal fungi to environmental change is an important step to maintaining forest productivity in the future. These predictions are currently limited by an incomplete understanding of the relative significance of environmental drivers in determining the community composition of ectomycorrhizal (ECM) fungi at large spatial scales. To identify patterns of community composition in ECM fungi along regional scale gradients of climate and nitrogen deposition in Scotland, fungal communities were analysed from 15 seminatural Scots pine (Pinus sylvestris L.) forests. Fungal taxa were identified by sequencing of the ITS rDNA region using fungal‐specific primers. Nonmetric multidimensional scaling was used to assess the significance of 16 climatic, pollutant and edaphic variables on community composition. Vector fitting showed that there was a strong influence of rainfall and soil moisture on community composition at the species level, and a smaller impact of temperature on the abundance of ectomycorrhizal exploration types. Nitrogen deposition was also found to be important in determining community composition, but only when the forest experiencing the highest deposition (9.8 kg N ha^?1 yr^?1) was included in the analysis. This finding supports previously published critical load estimates for ectomycorrhizal fungi of 5–10 kg N ha^?1 yr^?1. This work demonstrates that both climate and nitrogen deposition can drive gradients of fungal community composition at a regional scale.     

Uncertainties in the relationship between atmospheric nitrogen deposition and forest carbon sequestration

In a recent study, Magnani et al. report how atmospheric nitrogen deposition drives stand-lifetime net ecosystem productivity (NEP_av) for midlatitude forests, with an extremely high C to N response (725 kg C kg⁻¹ wet-deposited N for their European sites). We present here a re-analysis of these data, which suggests a much smaller C : N response for total N inputs. Accounting for dry, as well as wet N deposition reduces the C : N response to 177 : 1. However, if covariance with intersite climatological differences is accounted for, the actual C : N response in this dataset may be <70 : 1. We then use a model analysis of 22 European forest stands to simulate the findings of Magnani et al. Multisite regression of simulated NEP_av vs. total N deposition reproduces a high C : N response (149 : 1). However, once the effects of intersite climatological differences are accounted for, the value is again found to be much smaller, pointing to a real C : N response of about 50–75 : 1.     

Effects of nitrogen deposition on forest biodiversity

Humans have altered global and regional cycles of nitrogen (N) more than any other elements. Increasing N emissions to the atmosphere from accelerating industrialization and production and use of fertilizer N now make N deposition significant not only in densely populated regions of Europe and North America, but also in other parts of the world (e.g., Asia and Latin America). Increased atmospheric N deposition is known to be able to reduce biodiversity in natural and semi-natural ecosystems. It is suggested that N deposition will be the third greatest driver of biodiversity loss on the global scale in this century, after land use and climate change. Based on published study results, we reviewed the impacts of N deposition on forest biodiversity by emphasizing 3 aspects: (1) plant diversity, including arborous plants, understory plants and cryptogam plants; (2) soil microorganism diversity; (3) animal diversity, including underground soil fauna and aboveground herbivores. In general, it was found that N deposition could alter species diversity, and excessive N could reduce species diversity, such as richness and abundance, and even lose special species. We also identified specific mechanisms on how excessive N deposition affected forest biodiversity. Finally, we summarized the current status of research on N deposition in China and in other countries, and proposed potential research activities and recommendations.     

Effects of grass species and grass growth on atmospheric nitrogen deposition to a bog ecosystem surrounded by intensive agricultural land use

We applied a ¹⁵N dilution technique called “Integrated Total Nitrogen Input” (ITNI) to quantify annual atmospheric N input into a peatland surrounded by intensive agricultural practices over a 2-year period. Grass species and grass growth effects on atmospheric N deposition were investigated using Lolium multiflorum and Eriophorum vaginatum and different levels of added N resulting in increased biomass production. Plant biomass production was positively correlated with atmospheric N uptake (up to 102.7 mg N pot⁻¹) when using Lolium multiflorum. In contrast, atmospheric N deposition to Eriophorum vaginatum did not show a clear dependency to produced biomass and ranged from 81.9 to 138.2 mg N pot⁻¹. Both species revealed a relationship between atmospheric N input and total biomass N contents. Airborne N deposition varied from about 24 to 55 kg N ha⁻¹ yr⁻¹. Partitioning of airborne N within the monitor system differed such that most of the deposited N was found in roots of Eriophorum vaginatum while the highest share was allocated in aboveground biomass of Lolium multiflorum. Compared to other approaches determining atmospheric N deposition, ITNI showed highest airborne N input and an up to fivefold exceedance of the ecosystem-specific critical load of 5–10 kg N ha⁻¹ yr⁻¹.     

模拟氮沉降对温带阔叶红松林地土壤氮素净矿化的影响

以长白山阔叶红松混交林为研究对象,于2006—2008年原位模拟不同形态氮((NH4)2SO4、NH4Cl和KNO3)沉降水平(22.5和45kgN·hm-2·a-1),利用树脂芯法技术(resin-core incubation technique)测定了表层(有机层0～7cm)和土层(0～15cm)土壤氮素净矿化、净氨化和净硝化通量的季节和年际变化规律。同时,结合前人报道的有关林地碳、氮过程及其环境变化影响的结果,力求有效预估森林生态系统中氮素年矿化通量对大气氮沉降量和水热条件等因子变化的响应。结果表明,长白山阔叶红松林地土壤氮素年净矿化通量为1.2～19.8kgN·hm-2·a-1,2008年不同深度的土壤氮素年净矿化通量均显著高于2006和2007年(P<0.05)。随着模拟氮沉降量增加,土壤氮素净矿化通量也随之增加,尤其外源NH4+-N输入对净矿化通量的促进作用更为明显(P<0.05),但随着施肥年限的延长,这种促进作用逐渐减弱。与林地0～15cm土壤相比,氮沉降增加对0～7cm有机层氮素净氨化和净矿化通量的促进作用更为明显,尤其NH4Cl处理的促进作用更大。结合前人报道的野外原位观测结果,土壤氮素年净矿化通量随氮素沉降量的增加而增大,氮沉降量对不同区域森林土壤氮素净矿化通量的贡献率约为52%;氮沉降量(x1)和pH值(x2)可以解释区域森林土壤氮素年净矿化通量(y)变化的70%(y=0.54x1-18.38x2-109.55,R2=0.70,P<0.0001)。前人研究结果仅提供区域年均温度,未考虑积温的影响,这可能是造成年净矿化通量与温度无关的原因。今后的研究工作应该加强区域森林土壤积温观测,进而更加准确地预估森林土壤氮素的年净矿化通量。     

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。     

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Exploring cycad foliage as an archive of the isotopic composition of atmospheric nitrogen

Molecular nitrogen (N₂) constitutes the majority of Earth's modern atmosphere, contributing ~0.79 bar of partial pressure (pN₂). However, fluctuations in pN₂ may have occurred on 10⁷–10⁹ year timescales in Earth's past, perhaps altering the isotopic composition of atmospheric nitrogen. Here, we explore an archive that may record the isotopic composition of atmospheric N₂ in deep time: the foliage of cycads. Cycads are ancient gymnosperms that host symbiotic N₂‐fixing cyanobacteria in modified root structures known as coralloid roots. All extant species of cycads are known to host symbionts, suggesting that this N₂‐fixing capacity is perhaps ancestral, reaching back to the early history of cycads in the late Paleozoic. Therefore, if the process of microbial N₂ fixation records the δ¹⁵N value of atmospheric N₂ in cycad foliage, the fossil record of cycads may provide an archive of atmospheric δ¹⁵N values. To explore this potential proxy, we conducted a survey of wild cycads growing in a range of modern environments to determine whether cycad foliage reliably records the isotopic composition of atmospheric N₂. We find that neither biological nor environmental factors significantly influence the δ¹⁵N values of cycad foliage, suggesting that they provide a reasonably robust record of the δ¹⁵N of atmospheric N₂. Application of this proxy to the record of carbonaceous cycad fossils may not only help to constrain changes in atmospheric nitrogen isotope ratios since the late Paleozoic, but also could shed light on the antiquity of the N₂‐fixing symbiosis between cycads and cyanobacteria.     

氮沉降对森林土壤磷循环的影响

磷是生物体必需的大量元素之一,也是许多生态系统的主要限制因子。近年来,大气氮沉降日益加剧,已对森林生态系统磷循环产生了不可忽视的影响。关于氮沉降对生态系统磷循环的影响已开展了一系列的研究,然而尚缺少对其整体的认识。因此,通过收集国内外已发表的相关文章,从以下五个方面综述氮沉降对森林生态系统土壤磷循环的影响及其机理:1)阐述了森林生态系统土壤磷循环的概念;2)介绍了氮沉降对森林土壤磷循环影响的研究方法,包括长期定位模拟氮沉降法、自然氮沉降梯度法和同位素示踪法等;3)概述了氮沉降对森林生态系统土壤磷循环的影响。目前的研究结论趋向于认为长期氮沉降使森林土壤磷循环速率加快。长期氮输入易于使土壤中可溶性磷向非活性磷酸盐库迁移而难以被利用。因此,为了满足需求,土壤磷酸酶活性将增加以加速有机磷的矿化,从而加速磷素在土壤-植物-微生物之间的周转。4)探讨了氮沉降影响森林土壤磷循环的机制。氮沉降可通过改变土壤有机质的性质、微生物群落组成、磷酸酶活性以及阳离子的流动性等途径影响森林土壤磷循环;5)指出了当前研究存在的问题及未来的研究方向。     

川南天然常绿阔叶林土壤酶活性特征及其对模拟N沉降的响应

通过原位进行低氮(LN,50 kg N·hm-2·a-1)、中氮(MN,100 kg N·hm-2·a-1)和高氮(HN,150 kg N·hm-2·a-1)处理,研究了川南天然常绿阔叶林土壤酶活性特征及其对模拟N沉降的响应.结果表明,森林土壤过氧化氢酶、脲酶、纤维素酶和蔗糖酶的活性在垂直分布上均表现为0～10 cm土层高于10～20 cm土层;各种土壤酶活性的季节变化明显,但土壤酶活性还因取样时间和土层不同而异.其中,0～10 cm和10～20 cm土层中过氧化氢酶活性高峰均出现在秋季(92 d);0～10 cm土层中脲酶活性高峰出现在秋季,而10～20 cm土层中脲酶活性高峰出现在冬季(183 d);0～10 cm和10～20 cm土层中土壤纤维素酶活性高峰则出现在翌年春季(274 d);0～10 cm土层中蔗糖酶活性有两个高峰,分别出现在秋季和翌年春季;而10～20 cm土层中蔗糖酶活性只有一个高峰,出现在翌年春季.总体上,常绿阔叶林中不同土壤酶活性对N沉降的响应有所差异.其中,N沉降降低了常绿阔叶林土壤纤维素酶和过氧化氢酶活性,增加了土壤脲酶和蔗糖酶活性.不同浓度N处理间酶活性表现出了不同程度的差异性,但其变化的规律还不明显.     

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。     

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。     

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。