森林凋落物分解研究进展

森林凋落物是指森林生态系统内由生物组分产生,然后归还到林地表面的所有有机物质的总称。森林凋落物在促进森林生态系统正常的物质循环和养分平衡,维持生态系统功能中具有重要作用,其分解受多因素影响,且各因素之间相互交错。不同情况下,各因子的重要性可能不同。温度和湿度被认为是影响凋落物分解主要的气候因子。凋落物随着温度升高分解速率加快,增加土壤湿度对凋落物分解有积极作用。凋落物的化学性质中,C、N比和木质素含量被认为是最重要的指标。凋落物分解前期的分解速率受到养分含量、水溶性碳化合物和结构碳化合物含量的强烈影响,而后期则更多地受到木质索及纤维素／木质素比值的支配。土壤动物可以粉碎凋落物,土壤微生物也是促进凋落物分解的重要因素,人为活动也影响凋落物分解。N沉降、全球变暖和臭氧层破坏等全球变化对森林凋落物分解的影响已逐渐成为研究热点。未来凋落物分解的研究方向是统一研究方法,开展长期定位监测,加强对分解过程中有机碳含量和释放量的研究,以及N沉降对凋落物分解作用机理的研究。     

活性有机碳含量在凋落物分解过程中的作用

土壤凋落物的分解不仅是生态系统养分循环的重要环节,也是生态系统碳释放源之一。将呼伦贝尔森林草原过渡带的草原凋落物、白桦林凋落物、落松林凋落物分别添加在棕色针叶林土里进行恒温培养,探讨了不同凋落物类型有机碳分解速率差异及其影响因子。结果表明:不同凋落物的有机碳矿化速率和矿化累积总量在分解初期不一致,但由高到低的次序均为:草原凋落物→白桦林凋落物→落叶松林凋落物,40d的有机碳矿化累积量分别为76.53、47.42、20.56mg/g。这主要与凋落物的化学性质有关,主要决定于凋落物中易被微生物分解的热水溶性有机碳含量和易分解有机物含量,而与凋落物的总有机碳含量、全氮含量、w(C)/w(N)比等关系不明显。     

模拟氮沉降对温带森林凋落物分解的影响

森林凋落物的分解是生态系统养分循环的重要过程,以北京西山地带性植被栎树林（辽东栎：Quercus liaotungensis）为对象,主要研究温带森林植物凋落物分解对模拟氮沉降的响应,为更好地了解氮沉降对温带森林地区凋落物的分解过程提供参考.通过模拟氮沉降,研究不同形态氮（硝态氮、铵态氮和混合态氮）和不同水平氮沉降（对照0 kg·hm^-2·a^-1、低氮处理50 kg·hm^-2·a^-1 和高氮处理150 kg·hm^-2·a^-1）对凋落物分解的影响,在2 年的时间内调查分析了凋落物分解过程中质量损失动态和碳（C）、N 含量及w（C）/w（N）比值的变化.研究结果表明,氮沉降均使凋落物分解速率减缓,且随氮沉降剂量增加,凋落物分解速率相比对照分别减慢了9.88%（硝态氮低氮）、15.02%（硝态氮高氮）、11.46%（铵态氮低氮）、14.62%（铵态氮高氮）、13.04%（混合态氮低氮）和16.20%（混合态氮高氮）.且不同氮沉降类型、不同氮沉降水平间差异显著.不同形态、不同水平的氮沉降显著地增加了凋落物N 含量（P=0.061,P=0.087）,其中混合态氮沉降对凋落物中N 素含量增加最显著（P=0.044）.但在分解过程中,各处理均未对凋落物C 含量产生显著影响.不同水平的氮沉降显著降低了凋落物的w（C）/w（N）比值,而且不同类型不同水平氮沉降对凋落物w（C）/w（N）比值具有显著的交互作用（P=0.011）.综上所述,通过对模拟氮沉降后凋落物残留率等的变化分析,得出氮沉降对温带森林凋落物的分解产生了抑制作用.     

氮沉降下鼎湖山森林凋落物分解及与土壤动物的关系

研究了南亚热带3种森林生态系统凋落物在N沉降下的分解动态及其与土壤动物群落的关系。选取季风常绿阔叶林、针阔混交林和马尾松林建立野外模拟N沉降样地,实施四个处理组,对照(Control)、低氮(50kg·hm-2·a-1,LowN)、中氮(100kg·hm-2·a-1,MediujmN)和高氮处理(150kg·hm-2·a-1,HighN),利用凋落物网袋法,在18个月的时间内调查分析了凋落物分解过程及其中的土壤动物密度特征。研究结果表明,植被演替阶段对凋落物的分解速度存在影响,季风林凋落物降解速度显著性快于混交林和针叶林(P<0.05);18个月后,季风林各处理地凋落物残留率为0.05、0.14、0.13和0.17,混交林为0.64、0.56和0.62,针叶林为0.66、0.63和0.62。N沉降增加对凋落物分解存在一定影响。且这种影响与植被类型之间存在明显的交互作用。N沉降处理对季风林凋落物分解表现出了一定的抑制作用,而且这种差异随时间推移愈益明显,但在混交林和针叶林内,试验后期凋落物分解受到了N沉降处理的促进作用。在试验后期,尤其是12个月后,凋落物网袋土壤动物密度在不同林地和不同水平N处理下体现了差异化发展趋势。在季风林内,N处理地土壤动物密度受到了明显的抑制;在混交林和针叶林内,低N样地动物密度显示了相比对照样地的明显优势,但在较高强度的中N处理地无论在凋落物的降解速率还是在动物密度上都与对照样地没有明显差别。文章认为,N沉降处理所产生的影响可能受环境N饱和程度的调控。文章还提出,在凋落物分解进程中,土壤动物群落具有“后期进入”特征,这对于进一步准确分析森林凋落物分解进程及土壤动物的贡献有重要意义。     

华山松林凋落物养分释放及土壤生态化学计量特征对模拟氮沉降的短期响应

研究模拟氮(N)沉降下森林生态系统凋落物-土壤C/N/P化学计量特征,对探究在全球气候变化背景下森林生态系统物质循环内在机理具有重要科学意义。以滇中亚高山华山松林(Pinus armandii forest)为研究对象,采用尼龙网袋法于2018年2月—2019年1月在华山松林开展模拟N沉降下凋落叶、枝原位分解试验,分别设置4个N沉降水平:对照CK(N 0g·m~(-2)·a~(-1))、低氮LN(N 5 g·m~(-2)·a~(-1))、中氮MN(N 15 g·m~(-2)·a~(-1))和高氮HN(N 30 g·m~(-2)·a~(-1))。结果表明:华山松林凋落叶和枝C元素均为直接释放模式;凋落叶和枝N元素分别为淋溶-富集-释放和富集-释放模式;凋落叶和枝P元素分别为淋溶-富集-释放和富集-释放模式;凋落叶的C、N、P养分释放速率(40.71%、53.83%、47.06%)均高于凋落枝(20.98%、22.04%、13.15%);各N处理下,凋落叶和枝C释放速率均表现为LNMNCKHN;N沉降总体增加了凋落叶C、N含量,但对P含量无显著影响;N沉降显著降低了凋落叶ω_((C))/ω_((N))和ω_((N))/ω_((P))、凋落枝ω_((C))/ω_((N));凋落叶、枝N、P含量与土壤N、P含量密切相关,土壤P对凋落叶化学计量影响最大,土壤N对凋落枝化学计量影响最大,土壤C对凋落物化学计量影响最小。在短期内N沉降能抑制凋落物分解过程中C、N、P的释放,但对土壤化学计量特征无明显影响,滇中华山松林凋落物分解过程中的化学计量变化特征及养分释放的研究有助于了解森林生态系统对N沉降的响应机理,特别是土壤N、P对凋落物分解的影响将为后续研究的重点内容。     

丛枝菌根真菌对羊草凋落物降解作用的研究

丛枝菌根真菌（Abuscular mycorrhizal fungi）能够影响植物生长及养分含量,从而影响凋落物的降解。采用根袋的方法研究了接种两种丛枝真菌Glomus mosseae和Glomus claroideum对羊草（Leymus chinensis）地上部及根系凋落物降解的影响。结果表明,随时间的延长,凋落物的重量逐渐减少,凋落物氮、磷含量均表现出先下降后逐渐升高的趋势。接种对地上部凋落物的养分含量及降解速度未产生显著性影响,但显著降低了根系氮磷含量及降解系数。接种Glomus mosseae和Glomus claroideum羊草根系氮、磷含量均显著低于CK;接种与未接种相比羊草根系k值显著降低;根系C：N未接种处理显著低于接种处理。说明丛枝菌根真菌可能间接影响草原生态系统中有机物质的分解和养分释放。     

桂西北喀斯特次生林凋落物养分归还特征

以桂西北喀斯特次生林为研究对象,选择了自然恢复方式下处于同一演替序列的灌丛、藤剌灌丛、乔灌丛3个森林群落,应用网筐收集法于2007年9月至2008年8月定位观测和研究了各群落的凋落物量、组成特征、季节动态变化及其N、P、K含量.结果表明,3个次生林群落年均凋落物量范围为6 053.93 kg·hm-2(灌丛)～6 794.40 kg·hm-2(藤刺灌丛),凋落物以叶占明显优势,其年变化以单峰形式出现,峰值处在9月份;凋落物中主要养分元素的含量为N＞K＞P,而森林养分利用效率表现为P＞K＞N,灌丛、藤刺灌丛、乔灌丛的养分元素年归还总量分别为:95.41、106.09、80.62 kg·hm-2(N),6.58、6.64、5.06 kg·hm-2(P),18.63、19.66、14.90 kg-hm-2(K);群落地表凋落物层的凋落物现存量范围为2 835.23 kg·hm-2(藤刺灌丛)～3349.16 kg·hm-2(乔灌丛),凋落物的分解速率为1.82(灌丛)～2.37(藤刺灌丛),随着凋落物的分解养分元素的回归速度表现出K＞N＞P.同其他森林生态系统相比较,喀斯特次生林群落的凋落物量、养分归还量较大,养分回归较迅速,具有良好的自养能力和恢复潜力.     

鼎湖山马尾松林凋落物分解对凋落物输入变化的响应

为了了解我国南方森林常见的人为干扰（凋落物收取）活动对生态系统养分循环的影响,研究了鼎湖山马尾松林3种主要树种凋落物分解及其养分释放对凋落物输入量变化的响应。这3种树种分别为马尾松（Pinus massoniana）、荷木（Schimasuperba）和锥栗（Castanopsis chinensis）。凋落物输入量变化分别为凋落物去除（L-）、加倍（L＋）和对照（L）3种处理,每种处理25个重复。经过18个月的处理试验,凋落物分解速率及其养分释放随树种、分解阶段和凋落物处理不同而异。荷木、马尾松和锥栗分解物平均残留率分别为0.46±0.01、0.42±0.01、0.40±0.02,其中,荷木与锥栗、马尾松差异性显著。不同处理间的凋落物分解速率差异显著,加倍、对照和去除处理样地凋落物的平均残留率分别为0.51±0.08、0.53±0.09和0.55±0.08。凋落物加倍处理促进了凋落物分解过程中C的释放,而去除凋落物处理则抑制了N、P的释放。以上结果表明,凋落物收取活动不仅直接带走凋落物中的大量养分,而且抑制了凋落物分解及其养分释放。     

川西亚高山针叶林凋落物对土壤理化性质的影响

研究了川西地区亚高山人工云杉林及天然林凋落物变化及其对土壤理化性质的影响.结果表明30 a人工云杉林、40 a人工云杉林及次生林和原始林年凋落量分别为2.67×103 kg hm-2、4.38×103 kg hm-2、4.27×103 kg hm-2和4.77×103 kg hm-2,枯枝落叶层贮量分别为3.19×104 kg hm-2、3.64×104 kg hm-2、1.42×105 kg hm-2和1.45×105 kg hm-2,通过凋落物归还土壤的营养元素(N、P、K、Ca和Mg)的年归还总量依次为82.01 kg hm-2、129.04 kg hm-2、130.57 kg hm-2、170.55 kg hm-2,凋落物年失重率分别为24.35%、22.87%、36.96%和32.23%,人工林凋落物分解一半所需时间约为2.5 a,天然林约为1.6 a.各样地土壤含水量、孔隙度和养分含量大致表现为次生林≈原始林>30 a人工林>40 a人工林.森林年凋落量、枯枝落叶层贮量、养分归还量和年失重率与土壤自然含水率、有机质、N、P、K的含量呈正相关,与土壤容重呈负相关.人工云杉林生态功能的恢复滞后于次生林,凋落物分解缓慢是影响该地区土壤水分和养分状况的重要因素.人工云杉林进入旺盛生长期后,凋落量增加,养分归还量增大,此时期森林对土壤肥力有较高的补给潜力;但凋落物分解过缓,大量养分元素累积于枯枝落叶层,不能及时进入土壤,造成土壤理化性质状况较差.图1 表6 参18     

川西亚高山不同森林类型土壤酶活性对短期凋落物输入量变化的响应

凋落物输入是生态系统物质循环和能量流动的重要环节,在维持亚高山森林的"自肥"机制及生态系统结构和功能等方面具有不可替代的作用.为进一步了解川西亚高山森林土壤物质循环过程,以该地区典型森林为对象,通过进行凋落物输入控制试验,研究3种森林类型(阔叶林、针阔混交林和针叶林)不同时期(添加时间)凋落物输入变化对土壤碳(C)、氮(N)和磷(P)分解相关的酶活性及其化学计量比的影响.结果显示:3种森林类型中凋落物输入对土壤酶活性的影响不显著,而不同林型和凋落物输入时间则显著影响了土壤酶活性.凋落物处理1.5年时期,在凋落物输入处理中,阔叶林的β-葡萄糖苷酶和纤维二糖水解酶的酶活性显著高于针阔混交林(2.69和2.73倍)和针叶林(1.87和1.47倍);而在凋落物去除处理中,针叶林中的酸性磷酸酶活性则显著大于阔叶林(1.72倍)与针阔混交林(1.66倍).相关性分析表明,土壤含水量以及C、N含量是影响酸性磷酸酶、β-葡萄糖苷酶和纤维二糖水解酶变化的主要因素,而多酚氧化酶和过氧化物酶主要受到土壤pH以及土壤硝态氮含量的影响.根据生态酶化学计量理论,本研究的土壤酶化学计量碳氮比(C:NEEA)均低于全球平均值,而土壤酶化学计量碳磷比(C:PEEA)和土壤酶化学计量氮磷比(N:PEEA)均高于全球平均值,表明该区域N转化酶活性较高,处于相对N限制的区域.本研究表明在养分循环较慢的川西亚高山地区,短期凋落物的输入对土壤酶活性的影响较为缓慢,但不同凋落物组成(林型)对土壤酶活性的影响不同,关于土壤酶活性对于凋落物输入的响应还有待长期研究.(图4表3参49)     

Lignin and its effects on litter decomposition in forest ecosystems

Lignin is a major component of plant litter. In this review, we found lignin comprises a complex class of organic compounds whose concentration differs greatly both between and within plant species. There are many analytical methods for detecting the composition and structure of lignins. As lignins are enormously complex compounds, chemical assay is difficult and different methods vary with the results. Lignin plays a significant role in the carbon cycle, sequestering atmospheric carbon into the living tissues of woody perennial vegetation. It has also great effects on nitrogen dynamics of forest ecosystems as well as other ecological processes. Lignin is one of the most slowly decomposing components of dead vegetation, contributing a major fraction of the material that becomes humus as it decomposes. Lignin is highly correlated with decomposition of litter. Thus, there is evidence that the lignin concentration is a more influential factor than the other chemical concentrations, in determining the rate of leaf litter decomposition of different forest ecosystems. Although a great number of researchers have addressed lignin's role in litter decomposition, still there are many aspects of lignin biogeochemistry that are not known. This lack of information hinders complete amalgam of lignin effects on litter decomposition processes and dynamics of nutrient cycling.     

Tree species effects on decomposition and forest floor dynamics in a common garden

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.     

Long-term forest-floor litter dynamics in Canada's boreal forest: Comparison of two model formulations

Litter decomposition is a key component in ecosystem material cycling that determines (i) forest soil carbon (C) and nutrient content, (ii) release of carbon dioxide to the atmosphere, and (iii) generation and mass transfer of dissolved organic carbon from terrestrial to aquatic ecosystems. In this study, we provide simulations of long term forest-floor litter dynamics generated with both (i) an existing forest nutrient cycling and biomass growth model (ForNBM) with a single-pool formulation of forest-floor litter decomposition (Zhu et al., 2003. Ecol. Model. 169, 347-360), and (ii) a revised version of the model produced by substituting the single-pool formulation with a three-pool version of the formulation tested against data from litterbag experiments (FLDM; Zhang et al., 2010. Ecol. Model. 221, 1944-1953). This is done to determine the importance of subdividing the litter mass into categories of rates of decay (i.e., fast, slow, and very slow) on model accuracy. Forest-forest litter dynamics simulated with the two models are subsequently compared against field measurements collected at several northern jack pine (Pinus banksiana) stands along a southwest-northeast oriented transect (climate gradient) associated with the Boreal Forest Transect Case Study in northwest Canada. Initial comparison shows that the single-pool formulation underpredicts residual litter mass when forests are <65 years old, largely due to the improper treatment of the very slow decomposing litter component. This underprediction is resolved when the three-pool formulation is used. From a ecosystems-response point of view, the revised ForNBM (with the three-pool formulation) demonstrates that (i) forest-floor litter initially increases with forest growth and reaches a plateau once the forest matures; (ii) the forest floor stores more litter and C at the southern and warmer sites than at the northern sites; and (iii) in a similar climate regime, the forest floor stores more litter and C at productive than at nutrient-poor sites.     

Litter decomposition and nitrogen mineralization from an annual to a monthly model

The forest litter decomposition model (FLDM) described in this paper provides an important basis for assessing the impacts of forest management on seasonal stream water quality and export of dissolved organic carbon (DOC). By definition, models with annual time steps are unable to capture seasonal, within-year variation. In order to simulate seasonal variation in litter decomposition and DOC production and export, we have modified an existing annual FLDM to account for monthly dynamics of decomposition and residual mass in experimental litterbags placed in 21 different forests across Canada.The original annual FLDM was formulated with three main litter pools (fast, slow, and very slow decomposing litter) to address the fact that forest litter is naturally composed of a mixture of organic compounds that decompose at different rates. The annual FLDM was shown to provide better simulations than more complex models like CENTURY and SOMM.The revised monthly model retains the original structure of the annual FLDM, but separates litter decomposition from nitrogen (N) mineralization. In the model, monthly soil temperature, soil moisture, and mean January soil temperature are shown to be the most important controlling variables of within-year variation in decomposition. Use of the three variables in a process-based definition of litter decomposition is a significant departure from the empirical definition in the annual model. The revised model is shown to give similar calculations of residual mass and N concentration as the annual model (r² = 0.91, 0.78), despite producing very different timeseries of decomposition over six years. It is shown from a modelling perspective that (i) forest litter decomposition is independent of N mineralization, whereas N mineralization is dependent on litter decomposition, and (ii) mean January soil temperature defines litter decomposition in the summer because of winter-temperatures’ role in modifying forest-floor microorganism community composition and functioning in the following summer.     

Effects of visible light radiation on macrophyte litter degradation and nutrient release in water samples from a eutrophic shallow lake

Visible light is a major fraction of the solar spectrum; however, information on visible light radiation of macrophyte detritus is lacking. In this study, we conducted a microcosm experiment to assess the effects of visible light radiation on degradation of two litter species: Potamogeton malaianus (P. malaianus) and Phragmites australis (Ph. australis). This research represents an investigation of mass loss, microbial activity and nutrients released over a period of 168 days. Overall, we found that visible light radiation had significant effects on litter decomposition, but it did not affect the microbial activities which degrade cellulose and lignin. The decomposition rate order of the three components in P. malaianus and Ph. australis in treatments was: cellulose?>?hemicellulose?>?lignin. The visible light radiation mainly affected the degradation of lignin, which is the primary compound in litter susceptible to photodegradation. The exposure to visible light radiation up to 17.6?Wm^?2 stimulated the dissolved organic carbon release and reduced the molecular weight to less reactive. Meanwhile, no obvious difference in nutrient contents (TP, TN, NO₃–N, NO₂–N, and NH₃–N) was observed among different visible light intensities. The results of this study contribute to better understanding of the photochemical behaviour of macrophyte litter in shallow lakes.     

Impact of litter removal on soil characteristics under a Pinus caribaea stand

ABSTRACT

Forest productivity is dependent on soil quality, which in turn is related to litter; yet there is limited understanding of the relationships between litter and soil quality. The effects of litter removal treatment on tree growth and soil properties were examined in a Pinus caribaea stand with the aim of providing an understanding of consequences of litter removal on soil patterns. This knowledge is important for planning appropriate long-term forest management. Three pairs of 30 × 30 m² plots (each pair a control and treatment) were established in the center of a P. caribaea stand in April 2002. The controls were left undisturbed with an intact litter layer, while litter was removed monthly from treated plots. The diameter and height of the P. caribaea decreased and soil quality was degraded over the experimental period in litter-removed (treated) plots. Litter removal also produced a significant increase in soil bulk density at 0–20 and 20–40 cm depths. The capillary porosity, capillary moisture capacity, and natural water content in controls at 0–20 and 20–40 cm depths were significantly greater than treated plots. The non-capillary porosity in controls at the 0–20 cm depth was also significantly higher than treated plots. The organic carbon, total nitrogen (N), total potassium (K) and alkalized N content in each layer, and available P and exchangeable K at 0–20 and 40–60 cm depths in control plots were significantly greater than litter-removed plots. The numbers of bacteria at 0–20 and 40–60 cm depths and of fungi at 20–40 and 40–60 cm depths were higher in control than treated plots. The number of actinomyces and urease, catalase, and acid phosphatase activities in controls at each depth were significantly greater than litter-removed plots.