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素及纤维素/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。     

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。     

不同形态氮输入对湿地生态系统碳循环影响的研究进展

人类活动导致湿地生态系统氮负荷明显增加,引起生态系统碳循环过程发生诸多变化。外源氮输入对湿地生态系统土壤碳库稳定性的影响已成为当今国际研究的前沿问题之一。文章综述了不同形态氮素对湿地植物固碳潜势、土壤自养与异养呼吸速率的影响、土壤甲烷排放及不同形态氮与全球变暖对土壤有机碳及其组分矿化速率的交互作用的研究进展。研究表明,(1)植物对不同形态氮素的选择性吸收,会影响植物叶片的光合速率,改变植物的固碳潜势,影响植物根系的自养呼吸速率;同时,会影响凋落物归还量,改变植物对土壤的有机碳输入;此外,还可能影响凋落物的质量(如C/N),改变凋落物的分解速率,影响土壤异养呼吸速率。(2)各种形态氮输入对土壤p H产生不同的影响,改变土壤微生物及酶活性,影响有机碳的分解及土壤异养呼吸速率。(3)土壤有机碳组分对各种形态氮素的不同响应,也会改变土壤有机碳的矿化速率。(4)植物对不同形态氮素的选择性吸收,及各种形态氮输入对土壤p H产生的不同影响,会影响土壤中可利用C、N源的供应,改变土壤的酸碱环境及氧化还原电位,影响土壤CH4排放。(5)大气氮沉降与全球变暖同时影响土壤碳循环过程,但不同形态氮素与全球变暖对湿地土壤碳循环过程的交互作用研究仍较少见。迄今为止,氮沉降对湿地土壤碳库稳定性的影响效应仍存在很大的不确定性,仅有少量研究区分氮素形态对土壤碳库稳定性的影响,而关于湿地生态系统的研究鲜有报道。今后应着重区分不同形态氮素对湿地土壤碳库稳定性的影响机理研究,以便深入了解氮沉降与湿地土壤碳库稳定性之间的关系。     

季节性积雪下的高山草地凋落物分解

青藏高原高山草地的凋落物分解是其生物地化循环过程的重要一环.采用样方调查法估测两种典型地形条件下高山草地单位面积的年凋落物产量,分析定面积凋落物分解袋中的初始装袋量与凋落物分解率的关系,估算最佳初始凋落物装袋量.通过积雪期设置双面凋落物分解袋于浅雪、深雪、无雪及人工雪被处理下,比较不同的雪被处理下凋落物的分解率,同时测定对应的土壤温度及土壤微生物生物量碳氮含量,进而分析土壤微生物生物量碳氮与凋落物分解率之间的相关关系.研究发现:(1)两种典型样地内植物凋落物在自然状态下的年产生量均约为90 g/m~2;(2)非生长季中凋落物分解率与凋落物初始重量呈负线性相关,凋落物初始装袋量3 g是研究分解率的相对较佳重量;(3)随着积雪厚度的增加,非生长季土壤温度和凋落物分解率提高,同时也促进了土壤微生物生物量的积累,凋落物分解率和土壤微生物生物量碳、氮在深雪与无雪处理下分别达到最大值(分别为10.15%,156.37 mg/kg,75.89 mg/kg)和最小值(分别为3.07%,65.38mg/kg,20.17mg/kg).土壤微生物生物量碳、氮均与凋落物分解率呈显著相关性(P 0.05).综上所述,即便在冬季气温低于冰点的情况下,雪被的隔绝作用使得土壤微生物活动依然活跃进行;雪被变化既改变了土壤环境因子及凋落物分解率,也深刻影响着高山草地生态系统的结构与功能.     

湿地挺水植物凋落物立枯分解研究进展

在许多湿地生态系统中,挺水植物立枯常组成凋落物总量的主要部分,其分解过程研究对于正确评价湿地生态系统的碳收支状况具有重要意义。对立枯分解过程与传统研究中凋落物分解过程的差异,分解过程中的 CO2排放,立枯分解的研究方法,以及分解过程的影响因素进行了综述。一般认为,湿地挺水植物立枯与传统研究中凋落物所处环境的差异造成了分解过程的差异,其分解过程中产生 CO2是湿地温室气体排放的重要途径;凋落袋和红外气体分析仪是湿地挺水植物立枯分解研究的主要方法;真菌组成了立枯体上微生物的大部分,对于立枯分解具有重要作用;环境因子（温度、水分等）和“基质质量”是影响挺水植物立枯分解过程的重要因素。在全球变化背景下,气候变化因子会直接或间接影响凋落物立枯分解过程,引起对气候变化的反馈,因此,在全球范围内开展气候变化对于湿地生态系统挺水植物立枯分解影响的研究具有重要的意义,必将成为凋落物研究的一个重要方向。     

亚热带森林凋落物层土壤酶活性的季节动态

凋落物分解是森林生态系统养分循环中至关重要的过程。凋落物的分解与凋落物中的分解酶的活性直接相关。本研究调查了亚热带常绿阔叶林与针叶林中与土壤碳氮元素循环有关的β-糖苷酶(β-glucosidase)、内切纤维素酶(endocellulase)和几丁质酶(chitobiase)活性的季节变化规律。结果表明:在两个群落中,3种土壤酶活性与温度之间都有一定相关性,最高值均出现在温度较高的七月份;阔叶林中三种酶的活性在各个季节中大小顺序都为未分解凋落物层(L)>半分解层(F)>腐殖质层(H),而针叶林中则是半分解层(F)>未分解凋落物层(L)>腐殖质层(H);在每个取样时间点β-糖苷酶和几丁质酶的活性最高值都在阔叶林L层,显著高于针叶林L层,而内切纤维素酶的活性最高值是在针叶林H层。两林型下酶活性动态的差异,可能是导致两林型下凋落物层残存量差别的一个重要原因。     

内蒙古典型草原凋落物分解对不同草地利用方式的响应

以2005年设置的中德放牧控制试验为平台,研究了内蒙古典型草原群落凋落物的分解速率、失重率及其碳氮损失率在不同草地利用方式和放牧强度下的变化规律。草地利用方式包括:传统放牧、传统割草和放牧、割草轮换利用(混合利用)。放牧强度包括7个:0、1.5、3.0、4.5、6.0、7.5、9.0 sheep·hm~(-2)。结果表明:传统放牧和混合利用方式下,放牧都不同程度地加速了群落凋落物的分解和失重。在传统放牧平地系统中,放牧强度为7.5 sheep·hm~(-2)小区群落凋落物的分解速率和失重率均显著高于对照区,分别高出45.0%和40.0%;在混合利用平地系统中,放牧强度为6.0 sheep·hm~(-2)小区群落凋落物的分解速率显著高于对照区,高出35.0%;在混合利用坡地系统中,放牧强度为9.0 sheep·hm~(-2)小区群落凋落物的分解速率和失重率均显著高于对照区,分别高出82.4%和62.0%。群落凋落物碳损失率和氮损失率在传统放牧系统中,随放牧强度的增加呈现出相似的变化趋势,而在混合利用系统中,随放牧强度的增加呈现出相反的变化趋势,但是总体表现为,增加放牧强度会加速群落凋落物中碳氮的释放。高放牧强度下,群落凋落物的分解速率、失重率和碳损失率差异显著,表现为:混合利用传统割草传统放牧,这一试验结果在一定程度上验证了放牧优化假说。     

“非正常”凋落物对冰雪灾后南岭森林土壤有机碳的影响

2008年初,一场百年罕见的冰雪灾害袭击了我国南方地区,对南方19个省（区）的林区,相当中国森林面积的13%（1.86×107 hm2）的森林植被遭受整体性、毁灭性破坏,导致的大量“非正常”凋落物必将对森林土壤碳循环产生深远影响。本研究以广东乐昌杨东山十二度水省级自然保护区不同海拔梯度上受灾的常绿阔叶混交林3种森林群落为研究对象,采用去除冰雪灾害导致的“非正常”凋落物与受损林地进行对照试验,对冰雪灾害所导致“非正常”凋落物对土壤碳的影响进行了研究。研究结果表明：灾后3种森林群落中不同深度土壤的有机碳质量分数和储量持续增加,同一深度土壤有机碳质量分数和碳储量在不同年份之间差异显著。直至2011年“非正常”凋落物分解完毕时,10~50 cm的土壤有机碳质量分数与上一年相比有减少趋势;0~10 cm的土壤有机碳质量分数稍比2010年高,但变化差异不显著(p=0.36),说明在“非正常”凋落物分解结束后土壤有机碳质量分数有减小的趋势,致使土壤碳储量下降。去除“非正常”凋落物样地与对照样地相比,各土壤深度的土壤有机碳质量分数明显下降,年平均下降百分比例变化范围为12.14%~55.34%,0~10 cm和10~30 cm土壤的有机碳质量分数下降变化波动幅度要比30~50 cm大;不同深度土壤有机碳质量分数与凋落物有机碳失质量间具有显著的线性相关性。可见,冰雪灾害导致的“非正常”凋落物对土壤有机碳质量分数和储量都有十分显著的影响,“非正常”凋落物的分解输入能显著提高土壤有机碳质量分数和储量。     

川西地区7种常见彩叶树种凋落物分解

凋落物分解是森林土壤碳和养分的重要来源.彩叶林为川西地区重要的生态系统类型之一,为探索川西地区常见彩叶树种凋落物分解速率与养分释放,采用尼龙分解袋法研究常见彩叶物种葛罗枫(Acer grosseri)、落叶松(Larix kaempferi)、青榨槭(Acer davidii)、元宝枫(Acer truncatum)、黄栌(Cotinus coggygria)、红桦(Betula albosinensis)和花楸(Sorbuspohuashanensis)凋落物分解过程中的质量损失和养分释放特征.结果表明,分解两年后7种彩叶树种凋落物质量残留率从高到低依次为元宝枫(66.4%)、落叶松(64.1%)、红桦(63.7%)、青榨槭(59.6%)、黄栌(58.9%)、葛罗枫(50.8%)和花楸(50.1%);凋落物质量损失主要发生在第一年,第一年凋落物质量损失率显著高于第二年.两年分解过程中,7种彩叶树种凋落物碳浓度下降,碳表现为净释放.氮磷释放特征在物种间差异显著,其中,花楸凋落物氮磷均表现为净释放.相关性分析表明,初始养分(氮和磷)含量与分解速率呈正相关,而难分解物质(木质素和纤维素)含量与分解速率呈负相关.综上所述,川西彩叶树种凋落物分解和氮磷释放因树种和分解时期而不同,花楸凋落物分解和养分释放速率相对较快;研究结果可为川西彩叶林景观恢复重建的物种选择提供科学依据.(图3表3参41)     

Substrate concentration and enzyme allocation can affect rates of microbial decomposition

A large proportion of the world's carbon is stored as soil organic matter (SOM). However, the mechanisms regulating the stability of this SOM remain unclear. Recent work suggests that SOM may be stabilized by mechanisms other than chemical recalcitrance. Here, we show that the mineralization rate of starch, a plant polymer commonly found in litter and soil, is concentration dependent, such that its decomposition rate can be reduced by as much as 50% when composing less than approximately 10% of SOM. This pattern is largely driven by low activities of starch-degrading enzymes and low inducibility of enzyme production by microbial decomposers. The same pattern was not observed for cellulose and hemicellulose degradation, possibly because the enzymes targeting these substrates are expressed at constitutively high levels. Nevertheless, given the heterogeneous distribution of SOM constituents, our results suggest a novel low-concentration constraint on SOM decomposition that is independent of chemical recalcitrance. These results may help explain the stability of at least some SOM constituents, especially those that naturally exist in relatively low concentrations in the soil environment.     

Temperature sensitivity of the turnover times of soil organic matter in forests.

Soils represent the largest carbon pool in the terrestrial biosphere, and climate change might affect the main carbon fluxes associated with this pool. These fluxes are the production of aboveground litter and root litter, and decomposition of the soil organic matter (SOM) pool by soil microorganisms. Knowledge about the temperature sensitivity of the decomposition of different SOM fractions is crucial in order to understand how climate change might affect carbon storage in soils. In this study, the temperature sensitivity of the turnover times of three different SOM fractions (labile, intermediate, and stabilized) was investigated for 11 forest sites along a temperature gradient. Carbon-14 isotope analyses of the SOM fractions combined with a model provided estimates of their turnover times. The turnover times of the labile SOM fraction were not correlated with mean annual soil temperature. Therefore it was not possible to estimate temperature sensitivity for the labile SOM fraction. Given considerable evidence elsewhere for significant temperature sensitivities of labile SOM, lack of temperature sensitivity here most likely indicates limitations of the applied methodology for the labile SOM fraction. The turnover times of the intermediate and the stabilized SOM fractions were both correlated with mean annual soil temperatures. The temperature sensitivity of the stabilized SOM fraction was at least equal to that of the intermediate SOM fraction and possibly more than twice as high. A correction for confounding effects of soil acidity and clay content on the temperature sensitivities of the intermediate and stabilized SOM fractions was included in the analysis. The results as observed here for the three SOM fractions may have been influenced by (1) modeling assumptions for the estimation of SOM turnover times of leaf and needle longevities, constant annual carbon inputs, and steady-state SOM pools, (2) the occurrence of summer drought at some sites, (3) differences between sites in quality of the SOM fractions, or (4) the relatively small temperature range. Our results suggested that a 1 degree C increase in temperature could lead to decreases in turnover times of 4-11% and 8-16%, for the intermediate and stabilized SOM fractions, respectively.     

SOMPROF: A vertically explicit soil organic matter model

Most current soil organic matter (SOM) models represent the soil as a bulk without specification of the vertical distribution of SOM in the soil profile. However, the vertical SOM profile may be of great importance for soil carbon cycling, both on short (hours to years) time scale, due to interactions with the soil temperature and moisture profile, as well as on long (years to centuries) time scale because of depth-specific stabilization mechanisms of organic matter. It is likely that a representation of the SOM profile and surface organic layers in SOM models can improve predictions of the response of land surface fluxes to climate and environmental variability. Although models capable of simulating the vertical SOM profile exist, these were generally not developed for large scale predictive simulations and do not adequately represent surface organic horizons. We present SOMPROF, a vertically explicit SOM model, designed for implementation into large scale ecosystem and land surface models. The model dynamically simulates the vertical SOM profile and organic layer stocks based on mechanistic representations of bioturbation, liquid phase transport of organic matter, and vertical distribution of root litter input. We tested the model based on data from an old growth deciduous forest (Hainich) in Germany, and performed a sensitivity analysis of the transport parameters, and the effects of the vertical SOM distribution on temporal variation of heterotrophic respiration. Model results compare well with measured organic carbon profiles and stocks. SOMPROF is able to simulate a wide range of SOM profiles, using parameter values that are realistic compared to those found in previous studies. Results of the sensitivity analysis show that the vertical SOM distribution strongly affects temporal variation of heterotrophic respiration due to interactions with the soil temperature and moisture profile.     

Microbial nitrogen limitation increases decomposition

With anthropogenic nutrient inputs to ecosystems increasing globally, there are long-standing, fundamental questions about the role of nutrients in the decomposition of organic matter. We tested the effects of exogenous nitrogen and phosphorus inputs on litter decomposition across a broad suite of litter and soil types. In one experiment, C mineralization was compared across a wide array of plants individually added to a single soil, while in the second, C mineralization from a single substrate was compared across 50 soils. Counter to basic stoichiometric decomposition theory, low N availability can increase litter decomposition as microbes use labile substrates to acquire N from recalcitrant organic matter. This "microbial nitrogen mining" is consistently suppressed by high soil N supply or substrate N concentrations. There is no evidence for phosphorus mining as P fertilization increases short- and long-term mineralization. These results suggest that basic stoichiometric decomposition theory needs to be revised and ecosystem models restructured accordingly in order to predict ecosystem carbon storage responses to anthropogenic changes in nutrient availability.     

Nutrient regulation of organic matter decomposition in a tropical rain forest

Terrestrial biosphere-atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle's response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100-200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing >80% of their initial mass in <300 days. High organic matter solubility throughout decomposition combined with high rainfall support a model of litter-layer decomposition in these rain forests in which rapid mass loss in the litter layer is dominated by leaching of dissolved organic matter (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil.     

A process-based model of nitrogen cycling in forest plantations: Part II. Simulating growth and nitrogen mineralisation of Eucalyptus globulus plantations in south-western Australia

We applied a new version of the G’DAY ecosystem model to short-rotation plantations of Eucalyptus globulus growing under a Mediterranean climate in south-western Australia. The new version, that includes modified submodels for biomass production, water balance, litter and soil organic matter (SOM) decomposition, and soil inorganic N balance, was parameterised and applied to three experimental eucalypt sites (Mumballup, Darkan and Northcliffe) of contrasting productivity. With a common base set of parameter values, the model was able to correctly reproduce observed time series of soil water content, canopy leaf area index and stemwood data at the three sites. The model's ability to simulate soil N supply under forest plantations was tested by simulating N mineralisation at each of the three sites over the duration of the experiment (10 years). Simulated annual net N mineralisation in the litter and top 20 cm soil layer ranged from 50 to 170 kg N ha⁻¹ across the sites as a result of differences in rates of litter production, SOM and litter decomposition, and microbial N immobilisation and (re-)mineralisation. Simulations of annual soil N mineralisation were similar to measured rates over a 3-year period, except for an overestimation in 1 year at Mumballup and 2 years at Darkan. Model results indicated the importance of fine root production and turnover for N supply. As plantations age, supply of N to trees increasingly originates from litter decomposition, while the contribution from decomposition of SOM decreases. Although major soil feedbacks associated with litter production, decomposition and N availability are adequately integrated into G’DAY, further work is required in some aspects of the model, including the utility of the C-allocation submodel over a wide range of site conditions and silvicultural treatments.     

土地利用方式对土壤碳库影响的敏感性评价指标

综述了目前国内外在监测土地利用方式对土壤有机碳动态的影响时所采用的一些敏感性指标：微生物量碳和微生物商、CO2通量和qCO2、轻组有机质和颗粒态有机质、溶解态有机碳(DOC)。大量的研究表明，与土壤有机碳相比，微生物量碳库的周转率更大，周转时间更短，在土壤总有机碳变化可检测之前，土壤微生物部分的变化可能被检测到，是土壤碳动态的敏感性指标。轻组和颗粒态有机质是自然土壤肥力的决定因素，也是土地管理方式影响最明显的部分，对于准确评价土地利用变化对土壤碳过程的影响具有重要意义。CO2，通量和qCO2，可以综合反映土壤微生物的活性、利用土壤有机碳的效率及土壤中碳的代谢作用等，也是土壤碳动态的敏感性指标。DOC通量比全球植物和大气间碳交换量小1-2个数量级，所以生物圈碳平衡的很小变化会导致DOC的巨大变化，DOC浓度和通量是土壤温度和湿度变化的敏感指标。     

森林土壤有机层生化特性及其对气候变化的响应研究进展

森林土壤有机层是指累积在土壤表面未分解到完全分解的有机残余物质,在全球碳循环中具有十分重要的作用和地位.目前有关森林土壤有机层的生态研究主要集中于土壤有机层的凋落物储量、水土保持功能、生物多样性保育功能及其生化特性等,而有关其对气候变化响应的研究报道还相当少见,且已有的研究主要关于土壤有机层的碳源/汇动态等,有关森林土壤有机层生化特性对气候变化响应的研究还相对较少,这与其在全球气候变化中的作用和地位是极不相称的.过去10a中,有关土壤有机层生化特性对气候变化响应的研究主要包括土壤有机层的微生物数量、微生物生物量、呼吸作用、有机物质分解动态(凋落物分解)、酶活性等对环境变化的响应等方面.进一步的控制实验研究被认为是相当重要的.参51