全球气候变暖对凋落物分解的影响

凋落物分解作为生态系统核心过程,参与生态系统碳的周转与循环,影响生态系统碳的收支平衡,调控生态系统对全球气候变暖的反馈结果。全球气候变暖通过环境因素、凋落物数量和质量以及分解者3个方面,直接或间接地作用于凋落物分解过程,并进一步影响土壤养分周转和碳库动态。气候变暖可通过升高温度和改变实际蒸散量等环境因素直接作用于凋落物分解。气候变暖可引起植物物种短期内碳、氮和木质素等化学性质的改变以及群落中物种组成的长期变化从而改变凋落物质量。在凋落物分解过程中,土壤分解者亚系统作为主要生命组分(土壤动物和微生物)彼此相互作用、相互协调共同参与调节凋落物的分解过程。凋落物分解可以通过改变土壤微生物量、微生物活动和群落结构来加快微生物养分的固定或矿化,以形成新的养分利用模式来改变土壤有机质从而对气候变化做出响应。未来凋落物分解的研究方向应基于大尺度跨区域分解实验和长期实验,关注多个因子交互影响下,分解过程中碳、氮养分释放、地上/地下凋落物分解生物学过程与联系、分解者亚系统营养级联效应等方面。     

A global experiment suggests climate warming will not accelerate litter decomposition in streams but might reduce carbon sequestration

The decomposition of plant litter is one of the most important ecosystem processes in the biosphere and is particularly sensitive to climate warming. Aquatic ecosystems are well suited to studying warming effects on decomposition because the otherwise confounding influence of moisture is constant. By using a latitudinal temperature gradient in an unprecedented global experiment in streams, we found that climate warming will likely hasten microbial litter decomposition and produce an equivalent decline in detritivore-mediated decomposition rates. As a result, overall decomposition rates should remain unchanged. Nevertheless, the process would be profoundly altered, because the shift in importance from detritivores to microbes in warm climates would likely increase CO(2) production and decrease the generation and sequestration of recalcitrant organic particles. In view of recent estimates showing that inland waters are a significant component of the global carbon cycle, this implies consequences for global biogeochemistry and a possible positive climate feedback.     

Global negative vegetation feedback to climate warming responses of leaf litter decomposition rates in cold biomes

Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.     

Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes

Climate and litter quality have been identified as major drivers of litter decomposition at large spatial scales. However, the role played by soil fauna remains largely unknown, despite its importance for litter fragmentation and microbial activity. We synthesised litterbag studies to quantify the effect sizes of soil fauna on litter decomposition rates at the global and biome scales, and to assess how climate, litter quality and soil fauna interact to determine such rates. Soil fauna consistently enhanced litter decomposition at both global and biome scales (average increment ~ 27%). However, climate and litter quality differently modulated the effects of soil fauna on decomposition rates between biomes, from climate‐driven biomes to those where climate effects were mediated by changes in litter quality. Our results advocate for the inclusion of biome‐specific soil fauna effects on litter decomposition as a mean to reduce the unexplained variation in large‐scale decomposition models.     

亚热带森林凋落叶分解过程土壤节肢动物群落的变化特征

土壤动物群落结构和多样性可能随凋落物分解进程和基质质量的变化不断改变。为了解亚热带森林凋落叶分解过程中土壤节肢动物群落变化特征,以四川盆地亚热带森林麻栎（Quercus acutissima）和柳杉（Cryptomeria fortunei）凋落叶为对象,于2011-2015年采用分解袋法研究了2种凋落叶分解过程中土壤节肢动物组成、结构和多样性动态变化。整个研究期间,柳杉和麻栎凋落叶分解袋中共捕获土壤节肢动物3855只,分属于16目51科,且均以等节跳科和棘跳科为优势类群;麻栎凋落叶中土壤节肢动物的个体密度随分解进程呈现增加趋势,在分解的1079天达最高值后降低,而柳杉凋落叶则在分解的156天急剧增加后快速降低,2种凋落叶中土壤节肢动物类群数量具有相似的动态变化过程;2种凋落叶中土壤节肢动物总体以菌食性数量比例最高,腐食性最低,且随凋落叶分解进程,植食性土壤节肢动物占比明显下降,菌食性则上升;非度量多维尺度（NMDS）分析显示,2种凋落叶中土壤节肢动物群落组成具有显著差异,聚类分析表明,2种凋落叶土壤节肢动物群落结构相似性随分解进程不断降低。亚热带森林凋落叶分解过程中土壤节肢动物群落组成、结构和多样性受凋落叶类型影响。     

陆地生态系统凋落物分解对全球气候变暖的响应

陆地生态系统凋落物分解是全球碳收支的一个重要组成部分, 主要受气候、凋落物质量和土壤生物群落的综合控制。科学家们普遍认为全球气候变化将对陆地生态系统凋落物分解产生复杂而深远的影响。该文结合凋落物分解试验的常用方法——缩微试验、原位模拟实验和自然环境梯度实验, 归纳现有研究结果, 意在揭示全球气候变化对陆地生态系统凋落物分解的直接影响(温度对凋落物分解速率的影响)和间接影响(温度对凋落物质量、土壤微生物群落及植被型的影响)的普遍规律。各种研究方法都表明: 在水分条件理想的情况下, 温度升高往往能加快凋落物的分解速率; 原位模拟实验中, 凋落物分解速率因物种、增温方法和地理方位而异; 全球气候变化能改变凋落物质量, 但可能不会在短期内影响凋落物的分解速率; 凋落物质量和可分解性的种间差异远大于增温所引发的表型响应差异, 那么, 气候变化所引发的植物群落结构和物种组成的变化将对陆地生态系统凋落物分解产生更强烈的影响; 土壤生物群落如何响应全球气候变化, 进而怎样影响凋落物分解过程, 这些都还存在着极大的不确定性。     

陆地生态系统混合凋落物分解研究进展

凋落物分解在陆地生态系统养分循环与能量流动中具有重要作用,是碳、氮及其他重要矿质养分在生态系统生命组分间循环与平衡的核心生态过程。自然生态系统中,植物群落大多具有较高的物种丰富度和多样性,其混合凋落物在分解过程中也更有可能发生养分传递、化学抑制等种间互作,形成多样化的分解生境,多样性较高的分解者类群以及复杂的级联效应分解,这些因素和过程均对研究混合凋落物分解过程、揭示其内在机制形成了极大的挑战。从构成混合凋落物物种丰富度和多样性对分解生境、分解者多样性及其营养级联效应的影响等方面,综合阐述混合凋落物对陆地生态系统凋落物分解的影响,探讨生物多样性在凋落物分解中的作用。通过综述近些年的研究发现,有超过60%的混合凋落物对其分解速率的影响存在正向或负向的效应。养分含量有差异的凋落物混合分解过程中,分解者优先利用高质量凋落物,使低质量的凋落物反而具有了较高的养分有效性,引起低质量凋落物分解加快并最终使混合凋落物整体分解速率加快;而凋落物物种丰富度对土壤动物群落总多度有轻微的影响或几乎没有影响,但是对线虫和大型土壤动物的群落组成和多样性有显著影响,并随着分解阶段呈现一定动态变化;混合凋落物改变土壤微生物生存的理化环境,为微生物提供更多丰富的分解底物和养分,优化微生物种群数量和群落结构及其分泌酶的活性,并进一步促进了混合凋落物的分解。这些基于植物-土壤-分解者系统的动态分解过程的研究,表明混合凋落物分解作用不只是经由凋落物自身质量的改变,更会通过逐级影响分解者多样性水平而进一步改变分解速率和养分释放动态,说明生物多样性确实在一定程度上调控凋落物分解及其养分释放过程。     

湿地枯落物分解及其对全球变化的响应

综述了当前湿地枯落物分解及其对全球变化响应的研究动态。湿地枯落物分解研究已随研究方法的改进而不断深化;当前湿地枯落物分解过程研究主要集中在有机质组分和元素含量变化特征的探讨上;湿地枯落物分解同时受生物因素（即枯落物性质以及参与分解的异养微生物和土壤动物的种类、数量和活性等）和非生物因素（即枯落物分解过程的外部环境条件,包括气候条件、水分条件、酸碱度与盐分条件以及湿地沉积的行为与特征等）的制约;模型已成为湿地枯落物分解研究的重要手段,对其研究也在不断深化。还讨论了湿地枯落物分解对于全球变化的响应,指出全球变暖、大气CO2浓度上升、干湿沉降及其化学组成改变可能对枯落物分解产生的直接、间接和综合影响。最后,指出了当前该领域研究尚存在的问题以及今后亟需加强的几个研究方面。     

Global climate change and litter decomposition: more frequent fire slows decomposition and increases the functional importance of invertebrates

Of the many mechanisms by which global climate change may alter ecosystem processes perhaps the least known and insidious is altered disturbance regimes. We used a field-based experiment to examine the climate change scenario of more frequent fires with altered invertebrate assemblages on the decomposition of Eucalyptus leaves. Our design comprised three fire regimes [long-term fire exclusion (FE), long-term frequent burning (FB) and FE altered to FB (FEFB)] and two litter bag mesh sizes (8.0 and 0.2 mm) that either permitted or denied access to the leaf litter by most invertebrates. We found a significant interaction effect between fire regime and mesh size in losses of litter mass and net carbon (C). Compared with the regime of FE, with more frequent burning (FB and FEFB) the pace of decomposition was slowed by 41% (when access to litter by most invertebrates is not impeded). For the regime of FE, denying access to leaf litter by most invertebrates did not alter the pace of decomposition. Conversely, under regimes of frequently burning, restricting access to the litter by most invertebrates altered the pace of decomposition by 46%. Similar results were found for net C. For net losses of nitrogen (N), no interaction effects between fire regime and mesh size were detected, although both main effects were significant. Our results show that by modifying disturbance regimes such as fire frequency, global climate change has the potential to modify the mechanism by which ecosystems function. With more FB, decomposition is driven not only by fire regime induced changes in substrate quality and/or physiochemical conditions but through the interaction of disturbance regime with animal assemblages mediating ecosystem processes.     

山地森林表层土壤酶活性对短期增温及凋落物分解的响应

为了探究贡嘎山地区暗针叶林表层土壤酶活性对增温及凋落物分解的响应,采用开顶式生长箱(open top chamber)和加热电缆(OTC-cable)联合增温的方式模拟增温,同时定位监测实验小区地表空气、表层土壤温湿度的变化;不同类型(A:75%峨眉冷杉针叶+25%杜鹃荚蒾灌木叶凋落物,B:55%峨眉冷杉针叶+45%杜鹃荚蒾灌木叶凋落物)凋落物在模拟增温和自然对照条件下分解4年,研究土壤过氧化氢酶、多酚氧化酶和脲酶活性的变化。结果表明:增温使得地表空气和表层土壤温度分别增加了2.84℃和1.83℃;使得空气相对湿度和土壤含水量分别降低了5.27%和1.55%。针叶比例高会抑制凋落物分解,约13%;增温促进凋落物分解且对针叶比例高的促进作用更加明显,增幅均超过10%。增温总体上降低了土壤过氧化氢酶和脲酶活性,而对多酚氧化酶活性的影响表现为增大。针叶比例降低的凋落物分解对3种土壤酶活性的影响大致趋势是增大,幅度在15%以上。增温和凋落物类型之间的交互作用显著。随着土壤深度增加,土壤过氧化氢酶和多酚氧化酶活性增大,而脲酶活性降低。增温和不同类型凋落物分解对表层土壤酸碱性无显著作用。在土壤含水量较低的情况下,土壤水分对酶活性影响较大。贡嘎山峨眉冷杉林表层土壤酶对温度的敏感性不仅因酶类型、土壤深度而存在差异,也随增温时间、土壤水分条件而有所不同。     

四川盆地亚热带常绿阔叶林不同物候期凋落物分解与土壤动物群落结构的关系

为了解植物生长不同物候时期凋落物分解过程中土壤动物群落结构动态及其与凋落物分解的关系,以四川盆地亚热带常绿阔叶林典型人工林树种马尾松和柳杉,次生林树种香樟和麻栎凋落物为研究对象,采用凋落物分解袋试验研究,凋落物分解过程中土壤动物的群落特征。4种凋落物分解袋共获得土壤动物8047只,其中,柳杉(2341只)>香樟(2105只)>马尾松(2046只)>麻栎(1555只)。其中,秋末落叶期、萌动期和展叶期,马尾松凋落物袋中主要以捕食性土壤动物为优势类群,而后以菌食性土壤动物为主;香樟凋落物袋中除秋末落叶期和叶衰期以菌食性土壤动物为主要优势类群外,其他各时期均以捕食性土壤动物为主要优势类群;柳杉凋落物分解各时期均以菌食性土壤动物为主要优势类群;麻栎凋落物分解在前3个时期以菌食性为主,而后以植食性土壤动物为主要优势类群。相关分析表明,在秋末落叶期和萌动期土壤动物的个体密度主要和氮、磷含量及其格局密切相关,叶衰期主要和难分解组分木质素显著相关。除在秋末落叶期土壤动物对凋落物分解的贡献率与土壤动物的个体密度显著相关外,其余主要物候关键时期均与土壤动物的类群密度及其食性显著相关。     

中国温带和亚热带8个树种叶和吸收根协同分解

叶和细根(2mm)是森林生态系统的分解主体,二者是否协同分解,将极大影响所属植物在生态系统碳(C)循环中的物种效应。已有研究显示,叶和细根的分解关系具有极大的不确定性,认为很大程度上归因于细根内部具有高度的异质性,导致叶和细根在功能上不相似。为此,使用末梢1级根和细根根枝作为研究对象,它们在功能上同叶类似,称为吸收根。通过分解包法,分别在黑龙江帽儿山和广东鹤山,研究了2个阔叶树种和2个针叶树种(共8个树种)的叶和吸收根持续2a多的分解。结果发现,分解速率k(a~(-1),负指数模型)在8个树种整体分析时具有正相关关系(P0.05),在相同气候带或植物生活型水平上是否相关,受叶的分解环境及吸收根类型的影响;N剩余量整体上并不相关,亚热带树种的叶和细根根枝的N剩余量在分解1a后高度显著正相关,温带树种的叶和1级根的N剩余量在分解2a后显著高度正相关。本研究中,根-叶分解过程是否受控于相同或相关的凋落物性质是决定根-叶分解是否相关的重要原因,其中分解速率与酸溶组分正相关、与酸不溶组分负相关。比较已有研究,总结发现,根-叶分解关系受物种影响较大,暗示气候变化导致物种组成的改变将极大影响地上-地下关系,也因此影响生态系统C循环。     

Rates of litter decomposition in terrestrial ecosystems: global patterns and controlling factors

Aims We aim to construct a comprehensive global database of litter decomposition rate (k value) estimated by surface floor litterbags, and investigate the direct and indirect effects of impact factors such as geographic factors (latitude and altitude), climatic factors (mean annual tempePlrature, MAT; mean annual precipitation, MAP) and litter quality factors (the contents of N, P, K, Ca, Mg and C:N ratio, lignin:N ratio) on litter decomposition.Methods We compiled a large data set of litter decomposition rates (k values) from 110 research sites and conducted simple, multiple regression and path analyses to explore the relationship between the k values and impact factors at the global scale.Important findings The k values tended to decrease with latitude (LAT) and lignin content (LIGN) of litter but increased with temperature, precipitation and nutrient concentrations at the large spatial scale. Single factor such as climate, litter quality and geographic variable could not explain litter decomposition rates well. However, the combination of total nutrient (TN) elements and C:N accounted for 70.2% of the variation in the litter decomposition rates. The combination of LAT, MAT, C:N and TN accounted for 87.54% of the variation in the litter decomposition rates. These results indicate that litter quality is the most important direct regulator of litter decomposition at the global scale. This data synthesis revealed significant relationships between litter decomposition rates and the combination of climatic factor (MAT) and litter quality (C:N, TN). The global-scale empirical relationships developed here are useful for a better understanding and modeling of the effects of litter quality and climatic factors on litter decomposition rates.     

鼎湖山季风常绿阔叶林凋落叶分解与土壤动物群落动态和多样性

2001年12月~2002年12月,采用不同孔径分解凋落叶样袋法,对鼎湖山季风常绿阔叶林3类凋落叶的分解进行了研究,并对落叶分解过程中凋落叶袋内和袋下土样中的土壤动物群落和多样性进行了调查。结果表明,3种孔径袋内凋落叶的分解速率为大孔>中孔>微孔;混合凋落叶的分解速率大于单种凋落叶;蜱螨目在凋落叶分解的整个过程中相对数量都较高,弹尾目在凋落叶的分解过程中在凋落叶袋和土壤间移动,数量变化较大。凋落叶袋内大、中型土壤动物的个体数量在分解前期较多,中、小型土壤动物在分解的中期数量剧增;凋落叶袋内土壤动物的个体数量、密度以及多样性指数都随着落叶的分解而增加,9月最高;土壤样内则在分解的前期较高,以后逐渐降低。凋落叶的分解和土壤动物群落动态及多样性受凋落叶基质质量以及样地温度、降雨量等综合因素的影响。     

Chemistry and toughness predict leaf litter decomposition rates over a wide spectrum of functional types and taxa in central Argentina

Litter decomposition, a major determinant of ecosystem functioning, is strongly influenced by the litter quality of different species. We aimed at (1) relating interspecific variation in leaf litter decomposition rate to the functional types different species belong to; and (2) understanding the chemical and/or physical basis for such variation and its robustness to environmental factors. We selected 52 Angiosperms from a climatic gradient in central-western Argentina, representing the widest range of functional types and habitats published so far. Ten litter samples of each species were simultaneously buried for 9 weeks during the 1996 summer in an experimental decomposition bed. Decomposition rate was defined as the percentage of dry mass loss after incubation. Chemical litter quality was measured as carbon (C) content, nitrogen (N) content, and C-to-N ratio. Since tensile strength of litter and living leaves were strongly correlated, the latter was chosen as an indicator of physical litter quality. A subset of 15 species representing different functional types was also incubated in England for 15 weeks, following a similar experimental procedure. Litter C-to-N and leaf tensile strength of the leaves showed the strongest negative associations with decomposition rate, both at the species and at the functional-type level. Decomposition rates of the same species in Argentina and in England were strongly correlated. This reinforces previous evidence that species rankings in terms of litter decomposition rates are robust to methodological and environmental factors. This paper has shown new evidence of plant control over the turnover of organic matter through litter quality, and confirms, over a broad spectrum of functional types, general models of resource allocation. The strong correlations between leaf tensile strength – a trait that is easy and quick to measure in a large number of species – decomposition rate, and C-to-N ratio indicate that leaf tensile strength can be useful in linking plant quality to decomposition patterns at the ecosystem level. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of hydrologic changes on aboveground production and surface water chemistry in two boreal peatlands in Alberta: Implications for global warming

Aboveground net primary production (NPP) and surface water chemistryvariables were monitored in a lacustrine sedge fen and a bog for four years.There were no significant differences in precipitation, mean growing seasonannual temperature, and number of growing degree days from 1991 to 1994. Themean annual water levels in the lacustrine sedge fen differed significantly,whereas they were similar in the bog during these four years. We measured 15surface water variables in the lacustrine sedge fen and the bog, and foundthat only two correlated significantly with water level fluctuations. In thelacustrine sedge fen, calcium correlated positively (r²= 0.56) and nitrate correlated negatively (r² =0.20) with water levels. In the bog, potassium correlated positively(r² = 0.88) and total dissolved phosphorus correlatednegatively (r² = 0.62) with water levels. The remainingchemical variables showed no significant correlations with water levelfluctuations. Net primary production of the different vegetation strataappeared to respond to different environmental variables. In the lacustrinesedge fen, graminoid production was explained to a significant degree bywater levels (r² = 0.53), whereas shrub production wasexplained to a significant degree by surface water chemistry variables, suchas nitrate (r² = 0.74) and total phosphorus(r² = 0.22). In the bog, temperature was the onlyvariable that explained moss production to a significant degree(r² = 0.71), whereas ammonium explained graminoidproduction (r² = 0.66) and soluble reactive phosphorusexplained shrub production to significant degrees (r² =0.71). There are few direct data on the impact of climatic warming in borealwetlands, although paleoecological and 2×CO₂ model datahave provided some indications of past and possibly future changes invegetation composition, respectively. Our results suggest that thelacustrine sedge fen may succeed to a bog dominated by Sphagnum spp. andPicea mariana, whereas the bog may succeed to an upland-type forestecosystem. This revised version was published online in July 2006 with corrections to the Cover Date.