木兰围场典型落叶松-杨桦混交林生物量及固碳能力

以木兰林管局北沟林场内典型落叶松-杨桦混交林、落叶松人工林、白桦天然次生林、山杨天然次生林为研究对象,利用分层切割法和分层挖掘法对华北落叶松、白桦、山杨的生物量进行测定,并通过解析木进行了生长量的测定,从而建立生物量、生长量模型对林分的碳储量和固碳能力进行了估算。其研究结果表明:落叶松-杨桦混交林较落叶松人工林、白桦天然次生林、山杨天然次生林具有一定幅度的增产效益。落叶松-杨桦混交林中落叶松、白桦、山杨表现均优于各自的人工林或天然林,平均胸径分别高出6.7%、12.8%、4.1%,平均树高分别高出12.1%、1.4%、11.1%。落叶松-杨桦混交林中落叶松、白桦、山杨的固碳量增幅分别为29.74%、28.36%、34.52%;落叶松人工林固碳量增幅27.09%;白桦天然次生林固碳量增幅26.34%;山杨天然次生林固碳量增幅26.24%。落叶松-杨桦混交林中落叶松、白桦、山杨固碳量的增幅分别高于所对应树种的2.65%、2.02%、8.28%。     

北京东灵山优势种群树木死亡对空间格局与生境的影响

森林群落动态及其与环境因子关系研究一直是生态学的重要研究内容之一,树木死亡是森林群落动态的重要环节,环境因子是树木死亡的重要影响因素,因此,开展对树木死亡及其与环境相关性研究对分析森林群落动态至关重要。以东灵山暖温带落叶阔叶次生林20 hm~2动态监测样地为研究对象,对样地优势种群树木死亡前后(死亡前:活立木;死亡后,枯立木、倒木)种群数量特征、空间分布及生境相关性的变化关系进行研究,运用点格局、物种-生境相关性分析等方法对数据进行分析。结果:(1)树木死亡前,多度排序为辽东栎黑桦山杨华北落叶松;树木死亡后,多度排序为山杨黑桦辽东栎华北落叶松;树木死亡前后,随取样尺度增大,分布模式均由聚集向随机过渡。(2)种群与生境关联性分析表明:山杨、黑桦、华北落叶松活立木随机分布,辽东栎聚集于缓坡;山杨、辽东栎枯立木缓坡分布较多,黑桦随机,华北落叶松集中于山脊;山杨、黑桦倒木陡坡较多,辽东栎集中于山脊。(3)不同种群死亡前后的空间格局有明显的尺度依赖性,同时与物种自身生物学特性、密度制约、生境过滤的过程相关。     

山西太岳山森林演替过程中建群种的动态特征和种间关系

为解释太岳山白桦(Betula platyphylla)林向华北落叶松(Larix principis-rupprechtii)林演替的生态过程,该研究以不同演替阶段的调查样地数据为基础,通过空间代替时间构建演替的时间序列,即从白桦林演替至白桦+华北落叶松混交林再到华北落叶松林的过程;采用点格局分析法和Monte-Carlo拟合检验对演替过程中白桦和华北落叶松种群的分布格局及其相互关系进行研究。结果表明:(1)山西太岳山白桦种群龄级结构呈"纺锤"型,种群呈衰退的趋势;白桦种群逐渐被华北落叶松(龄级结构呈"金字塔"的增长型)种群所取代。(2)从白桦林演替至华北落叶松林的过程中,白桦种群由集群分布趋向于随机分布,驱动力主要是华北落叶松侵入扩散而形成的种间排斥;华北落叶松集群尺度逐步增大,驱动力主要是种群拓殖和种内竞争。(3)白桦和华北落叶松二者的种间关系,在白桦林阶段无显著相关,在混交林阶段由于排斥呈现显著负相关关系,华北落叶松林阶段二者无明显相关,二者种间关系的变化主要来自于群落剩余资源驱动下的种内种间竞争。     

关帝山次生杨桦林种群结构与立木的空间点格局

选择关帝山典型天然次生杨桦林样地进行种群结构分析,并运用空间点格局分析方法O-ring统计系统分析了次生杨桦林群落内不同尺度下种群空间分布格局及种间关联性.结果表明,在个体组成上,演替先锋树种白桦(Betula platyphylla)、红桦(B.albo-sinensis)和山杨(Populus davidiana)是目前次生杨桦林样地的优势种.但从种群径级结构看,白桦、红桦和山杨天然更新不良,而云杉(Picea spp.)和华北落叶松(Larix principis-rupprechtii)幼苗呈现良好的更新状态.随着演替的进行,云杉、落叶松将逐渐进入林冠层并取代白桦、红桦和山杨成为该森林的优势种;从种群空间分布格局来看,红桦、白桦表现为小尺度上明显聚集,华北落叶松、皂柳(Salix wallichiana)表现为小尺度上强聚集性分布格局.种群空间分布格局受空间尺度、群落结构及种群径级结构综合作用的影响;空间关联性分析表明,红桦与华北落叶松、红桦与皂柳、皂柳与华北落叶松在小尺度上正相关,其它树种间没有表现出明显关联性.     

树种相互作用、林分密度和树木大小对华北落叶松生产力的影响

本研究以塞罕坝地区华北落叶松纯林、白桦纯林和华北落叶松与白桦混交林为对象,每种林分类型设置2个林分密度(200～340和880～1100株·hm^-2)。基于树木大小分层取样,采集树芯样本668条。运用线性混合模型,分析了树种相互作用、林分密度和树木大小对华北落叶松生产力的影响。结果表明: 华北落叶松断面积生长量不同程度地受竞争、胸径、年龄和邻体密度的影响;在林分密度较高的混交林中,由于白桦对华北落叶松生长的促进作用,落叶松生产力发生明显的增益效应;在林分密度较低的混交林中,落叶松和白桦之间不发生相互作用,2个树种的生产力均低于相应的纯林;种内竞争是影响华北落叶松生产力的主要因素;树木大小对华北落叶松生产力产生积极的影响,但影响程度因林分密度和树种组成而异。适当增加林分密度和选择白桦作为混交树种可以提高华北落叶松生产力。     

山西太岳山好地方典型植被类型土壤理化特征

以山西太岳山好地方林场4种植被类型(草甸、灌丛、华北落叶松人工林、华北落叶松-白桦混交林)0~60 cm土层土壤为研究对象,采取野外定点取样和实验室分析的方法,研究不同植被类型土壤理化性质特征。结果表明:土壤容重和p H值随土层深度的增加而增加;土壤含水量随土层深度增加而减小;不同植被类型间,土壤含水量大小排序为华北落叶松-白桦混交林草甸华北落叶松人工林灌丛;容重为灌丛华北落叶松-白桦混交林华北落叶松人工林草甸;p H值变化规律为华北落叶松-白桦混交林华北落叶松人工林灌丛草甸;土壤有机质、全氮含量随土层深度增加而减小,各土层之间差异显著;全磷和全钾含量各土层间差异不显著;不同植被类型间,草甸土壤有机质、全氮、全磷含量都高于其他3种植被类型,灌丛土壤全钾含量高于其他3种植被类型;土壤有机质与全氮呈极显著正相关;容重与含水量、有机质、全氮呈极显著负相关;p H值与有机质、全氮、全磷呈极显著负相关。     

大兴安岭呼中地区白卡鲁山植物群落结构及其多样性研究

通过沿海拔梯度系统调查,运用TWINSPAN分类方法,划分出大兴安岭地区白卡鲁山的主要植物群落类型并分析了其结构特征。利用物种丰富度和α多样性等指标,研究了乔木层、灌木层和草本层植物的多样性特征及其随海拔的变化趋势。结果显示：(1)白卡鲁山乔木层在海拔。700-1290m范围内,可分为4种主要群落类型：兴安落叶松疏林、兴安落叶松-白桦混交林、兴安落叶松-白桦-樟子松混交林、兴安落叶松纯林群落;(2)同一植物群落的结构特征比较相似,除兴安落叶松疏林外,其他三种森林类型的平均胸径和平均树高随海拔升高有逐渐增大的趋势;(3)乔木层的直径和树高分布清晰地反映了群落的水平和垂直结构。直径分布均呈现倒“J”型,表明森林的自我更新状况良好;而树高分布主要以双峰分布和近似对称分布为主;(4)乔木层的物种丰富度和α多样性在兴安落叶松纯林中最小,在兴安落叶松-白桦混交林和兴安落叶松-白桦-樟子松混交林群落次之,而在兴安落叶松疏林群落相对较高。各群落类型的灌木层和草本层植物多样性没有明显的差异。乔木层、灌木层和草本层的物种多样性随海拔梯度均没有表现出明显的变化趋势。     

环境因子对山西太岳山典型森林类型物种多样性及其功能多样性的影响

该研究采用典型样地法,调查群落内物种分布并测量植物功能性状(叶面积和植株高度),对山西太岳山不同坡位华北落叶松-白桦混交林以及辽东栎次生林物种多样性及其功能多样性进行比较分析,探究环境因子对不同群落层次(乔木、灌木、草本)物种多样性及其功能多样性的影响机制,以及环境因子与群落构建之间的联系,为森林生态系统多样性研究以及经营管理提供理论依据。结果显示:(1)华北落叶松-白桦混交林的物种分布更加均匀,物种多样性和功能多样性(乔木层)均显著高于辽东栎次生林。(2)华北落叶松-白桦混交林乔木层功能均匀度与功能分散指数显著高于辽东栎次生林,但灌木草本层低于辽东栎次生林。(3)不同群落层次的物种多样性与功能多样性均呈正相关关系,影响物种分布和性状分布的环境因子存在差异,物种多样性受多种环境因子的综合影响,而单个环境因子对功能多样性影响较大,环境解释力与林分类型和群落层次相关。(4)乔木层物种多样性主要受土壤pH、冠层结构(MLA、林分开度)以及光照影响,灌木层物种多样性与土壤pH和MLA密切相关,林下总辐射、土壤养分(SOC、STN)、土壤相对含水率是影响草本层物种分布的主要环境因子;冠层结构(MLA、林分开度)是影响乔木层功能多样性最主要的环境因子,土壤pH和坡位分别是华北落叶松-白桦混交林和辽东栎次生林灌木层功能多样性的主要影响因子,影响草本层功能多样性的主要环境因子是土壤相对含水率与LAI。研究表明,在垂直分层的森林生态系统中,不同群落层次竞争的主要环境资源存在差异,乔木层通过改变冠层结构和林内环境限制林下物种分布和性状分布。     

大兴安岭新林林业局3种林分类型天然更新幼苗幼树的空间分布格局

基于大兴安岭新林林业局落叶松林、白桦林和混交林3种主要林分类型16块样地的调查数据,依照幼苗幼树的株高划分为3个等级: Ⅰ级, 株高≤60 cm; Ⅱ级, 60 cm＜株高≤200 cm; Ⅲ级, 株高＞200 cm且胸径＜5 cm.在4种取样尺度下,采用负二项指数等7种聚集度指数,判定幼苗幼树的分布格局及格局强度,分析格局变化和格局规模.结果表明: 除落叶松林的落叶松幼苗幼树Ⅱ级最多,且随等级升高幼苗幼树密度先升高后降低外,白桦林和混交林的落叶松以及3种林分的白桦幼苗幼树Ⅲ级最多,而Ⅰ级最少,随等级升高,幼苗幼树密度均升高.落叶松幼苗幼树在3种林分4种取样尺度下均呈聚集分布.白桦幼苗幼树在白桦林中除在10 m×20 m尺度为随机分布外,均呈聚集分布.除落叶松幼苗幼树在混交林中5 m×10 m尺度时聚集强度最大外,落叶松在其他林分以及白桦在3种林分均在5 m×5 m尺度下聚集强度最大.在落叶松林,落叶松和白桦幼苗幼树的聚集强度随等级增大而减小,其中白桦由聚集分布变为随机或均匀分布;在白桦林和混交林中,落叶松和白桦幼苗幼树的聚集强度随等级升高不断增大,由均匀分布变为聚集分布.在3种林分类型中,不同更新等级落叶松和白桦幼苗幼树种群的格局规模均为25 m².落叶松和白桦幼苗幼树种群格局强度在不同林分类型、不同更新等级中差异明显.     

大兴安岭中部天然次生林种子雨动态

本研究于2018—2019年对大兴安岭中部白桦林、针叶林和针阔混交林3种林型的种子雨落种量进行了动态监测,并对3种林型主要树种的种子雨季节动态、落叶动态、种子雨千粒重、种子雨年际变化和种子雨空间格局进行了分析。结果表明: 各林型中兴安落叶松种子雨和白桦种子雨呈现明显的单峰型分布。针阔树种(落叶松、樟子松、云杉、白桦、山杨)的落叶量呈现出明显的季节动态,各林型落叶量大多在9月中上旬达到高峰。在针阔混交林和针叶林中,处于高峰期的兴安落叶松种子雨千粒重明显大于初始期和末尾期的兴安落叶松种子雨千粒重,3种林型下白桦千粒重在季节上未表现出明显的差异。兴安落叶松和白桦的种子雨均呈现出明显的年际变化,2018年为种子散种量的丰年,2019年为歉年。两年时间内,所有种子雨的空间格局在总体上均表现为聚集分布,种子雨和幼苗幼树在空间分布格局上存在一致性。     

兴安落叶松分枝格局的分形特征

对于树木分枝格局分形特征的定量描述,可以加深对树木生长过程的理解。本文采用分形几何学对兴安落叶松(Larix gmelini)的分枝格局进行研究,结果表明1)兴安落叶松分枝格局是一种分形结构,存在自相似性。2)兴安落叶松分枝格局的分形维数介于1.4~1.7之间,揭示了它的结构复杂性程度和占据生态空间、利用生态空间的能力。分形维数在树木光合作用及生长发育研究中是一个有用的参数。     

Biomass production, relative competitive ability and water use efficiency of two dominant species in semiarid Loess Plateau under different water supply and fertilization treatments

Water stress and nutrient deficiency are considered to be the main environmental factors limiting plant growth and species interaction in semiarid regions. However, less is known about the interactive effects of soil water, nitrogen and phosphorus on native species growth and relative competitive ability. A replacement series design method was used with 12 mixed plants of Bothriochloa ischaemum and Lespedeza davurica grown in a pot experiment under three water regimes and four fertility treatments. Intercropping systems were assessed on the basis of indices such as biomass production and allocation, relative competitive ability, aggressivity, relative yield total and water use efficiency (WUE). Water stress decreased significantly the total biomass production for each species, either in monoculture or in mixtures. N, P, or NP application can significantly improve biomass production of the two species in their mixtures. There was no obvious change trend in root/shoot ratio of B. ischaemum or L. davurica in different mixture proportions. Relative yield total (RYT) values ranged from 0.98 to 1.39. Aggressivity values of B. ischaemum to L. davurica were positive in all water regimes and fertilizations, implying that B. ischaemum was the dominant species. Relative competition intensity values of B. ischaemum (i.e., RCI_B) were less than zero, while greater than zero for L. davurica (i.e., RCI_L), indicating that the effects of intraspecific competition with L. davurica were stronger for B. ischaemum, and the opposite for L. davurica. WUE increased gradually as the proportion of B. ischaemum increased in mixtures, and a 10:2 B. ischaemum:L. davurica mixture proportion had significantly higher WUE. Results suggest that it is advantageous to grow the two species together to maximize biomass production and the recommended mixture ratio was 10:2 of B. ischaemum to L. davurica because it gave higher RYT and significantly higher WUE under conditions of water deficit.     

Biomass production and relative competitiveness of a C3 legume and a C4 grass co-dominant in the semiarid Loess Plateau of China

Bothriochloa ischaemum L. and Lespedeza davurica (Laxm.) Schindl. are two co-dominant species of great importance in reducing soil and water loss and maintaining the distinctive natural scenery of the semiarid Loess Plateau of China. Our aim was to determine the growth and interspecific competition between these species under water stress to facilitate the prediction of community succession and guide the selection of appropriate methods of conservation and use in the area. A pot experiment was designed to investigate the effects of water stress and competition on biomass production and allocation, relative competitive ability and water use efficiency of the two species. Bothriochloa ischaemum (a C₄ perennial herbaceous grass) was planted in the same pot with L. davurica (a C₃ perennial leguminous subshrub) at density ratios of 12:0, 10:2, 8:4, 6:6, 4:8, 2:10, and 0:12. The response of the species to their mutual presence at the different ratios was evaluated at three levels of soil moisture (80%?±?5% field capacity, FC (HW), 60%?±?5% FC (MW) and 40%?±?5% FC (LW)). Indices of aggressivity (A), competitive ratio (CR) and relative yield totals (RYTs) were calculated from the dry shoot, root and total weight data. Water stress decreased the biomass production by both species whether in monoculture or mixture, but B. ischaemum was more sensitive to water deficit. Across moisture levels, the growth of L. davurica was more adversely affected by mixed planting. Bothriochloa ischaemum had significantly (P?<?0.05) smaller root:shoot ratios than L. davurica and the root mass of both species tended to increase relative to shoot mass as soil water deficit increased. The aggressivity (A), competitive ratio (CR) and relative yield totals (RYTs) of B. ischaemum were positive across treatments. Bothriochloa ischaemum had much higher CR under each water treatment, but water stress considerably reduced its relative CR while increasing that of L. davurica. The RYT values of the two species indicated some degree of resource complimentarity under both water sufficient and deficit conditions. Our results suggest that it is advantageous to grow the two species together to maximize biomass production. We recommend a mixture ratio of 8:4 of B. ischaemum to L. davurica because it gave significantly higher RYT and transpiration water use efficiency under deficit water conditions.     

Shovelomics: high throughput phenotyping of maize (Zea mays L.) root architecture in the field

We present a method to visually score 10 root architectural traits of the root crown of an adult maize plant in the field in a few minutes. Phenotypic profiling of three recombinant inbred line (RIL) populations of maize (Zea mays L.; B73xMo17, Oh43xW64a, Ny821xH99) was conducted in 2008 in a silt loam soil in Pennsylvania and in a sandy soil in Wisconsin, and again in 2009 in Pennsylvania. Numbers, angles and branching pattern of crown and brace roots were assessed visually at flowering. Depending on the soil type in which plants were grown, sample processing took from three (sand) to 8 min (silt-loam). Visual measurement of the root crown required 2 min per sample irrespective of the environment. Visual scoring of root crowns gave a reliable estimation of values for root architectural traits as indicated by high correlations between measured and visually scored trait values for numbers (r ²?=?0.46?C0.97), angles (r ²?=?0.66?C0.76), and branching (r ²?=?0.54?C0.88) of brace and crown roots. Based on the visual evaluation of root crown traits it was possible to discriminate between populations. RILs derived from the cross NY821 x H99 generally had the greatest number of roots, the highest branching density and the most shallow root angles, while inbred lines from the cross between OH43 x W64a generally had the steepest root angles. The ranking of genotypes remained the same across environments, emphasizing the suitability of the method to evaluate genotypes across environments. Scoring of brace roots was better correlated with the actual measurements compared to crown roots. The visual evaluation of root architecture will be a valuable tool in tailoring crop root systems to specific environments.     

太白红杉顶芽动态及其对分枝格局的影响

应用计盒维数方法比较研究了与太白红杉顶芽动态相关的不同分枝格局对空间占据能力的差异.结果表明,Ⅰ型分枝有利于对所拥有的空间进行有效填充和利用,Ⅱ型分枝有利于扩展枝条所占的面积,Ⅲ型分枝既扩展了面积,又实现了对所拥有空间的有效填充和利用.总的来讲,顶芽的死亡使分枝对空间的占据能力和对光能利用的效率有所提高,同时对植株冠型的调整也有重要作用.太白红杉枝条顶芽的死亡是外界严酷环境条件的被迫"产物".     

Biomass allocation, relative competitive ability and water use efficiency of two dominant species in semiarid Loess Plateau under water stress

A better understanding of the growth and interspecific competition of native dominant species under water stress should aid in prediction of succession in plant communities. In addition, such research would guide the selection of appropriate conservation and agricultural utilization of plants in semiarid environments that have not been very well characterized. Biomass production and allocation, relative competitive ability and water use efficiency of one C₄ herbaceous grass (Bothriochloa ischaemum) and one C₃ leguminous subshrub (Lespedeza davurica), both important species from the semiarid Loess Plateau of China, were investigated in a pot-cultivation experiment. The experiment was conducted using a replacement series design in which B. ischaemum and L. davurica were grown with twelve plants per pot, in seven combinations of the two species (12:0, 10:2, 8:4, 6:6, 4:8, 2:10, and 0:12). Three levels of water treatments included sufficient water supply (HW), moderate water stress (MW) and severe water stress (LW). These treatments were applied after seedling establishment and remained until the end of the experiment. Biomass production and its partitioning, and transpiration water use efficiency (TWUE) were determined at the end of the experiment. Interspecific competitive indices (competitive ratio (CR), aggressiveness (A) and relative yield total (RYT)) were calculated from the dry weight for shoots, roots and total biomass. Water stress decreased biomass production of both species in monoculture and mixture. The growth of L. davurica was restrained in their mixtures for each water treatment. L. davurica had significantly (P < 0.05) greater root:shoot allocation than B. ischaemum for each water treatment and proportion within the replacement series. Aggressiveness (A) values for B. ischaemum with respect to L. davurica were negative only at the proportions of B. ischaemum to L. davurica being 8:4 and 10:2 in LW treatment. B. ischaemum had a significantly (P < 0.05) higher CR value under each water treatment, and water stress considerably reduced its relative CR while increased that of L. davurica. RYT values of the two species indicated some degree of resource complimentarity under both water sufficient and deficit conditions. The results suggest that it is advantageous for growing the two species together to maximize biomass production, and the suggested ratio was 10:2 of B. ischaemum to L. davurica because of significantly higher (P < 0.05) RYT and TWUE under low water availability condition.     

Soil microbial properties under different vegetation types on Mountain Han

This study investigated the influence of broadleaf and conifer vegetation on soil microbial communities in a distinct vertical distribution belt in Northeast China.Soil samples were taken at 0-5,5-10 and 10-20 cm depths from four vegetation types at different altitudes,which were characterized by poplar(Populus davidiana)(1250-1300 m),poplar(P.davidiana) mixed with birch(Betula platyphylla)(1370-1550 m),birch(B.platyphylla)(1550-1720 m),and larch(Larix principis-rupprechtii)(1840-1890 m).Microbial biomass and community structure were determined using the fumigation-extraction method and phospholipid fatty acid(PLFA) analysis,and soil fungal community level physiological profiles(CLPP) were characterized using Biolog FF Microplates.It was found that soil properties,especially soil organic carbon and water content,contributed significantly to the variations in soil microbes.With increasing soil depth,the soil microbial biomass,fungal biomass,and fungal catabolic ability diminished;however,the ratio of fungi to bacteria increased.The fungal ratio was higher under larch forests compared to that under poplar,birch,and their mixed forests,although the soil microbial biomass was lower.The direct contribution of vegetation types to the soil microbial community variation was 12%.If the indirect contribution through soil organic carbon was included,variations in the vegetation type had substantial influences on soil microbial composition and diversity.     

六盘山典型森林伴随降水的总有机碳(TOC)通量变化特征

在六盘山香水河小流域,选择6种典型森林样地,测定了2011年生长季的大气降水、穿透水、干流、枯落物渗漏水和主根系层(0—30 cm深)土壤渗漏水的总有机碳(TOC)浓度及其相应的通量变化。结果表明,在降水转化为由穿透雨和干流组成的林下降水中,所有样地的TOC浓度都不同程度地增大;虽然林冠截持使林下降水减小,但因雨水淋洗和与林冠发生碳交换,各样地林下降水携带的生长季TOC通量(kg/hm2)(华北落叶松人工林132.28、华山松次生林106.56、油松人工林94.10、灌木林79.49、桦木林66.52、辽东栎次生林63.01)都比林外降水(53.17)不同程度地明显增大,整体看来,林冠的TOC淋出作用在针叶林很大,在阔叶林较弱。在6种森林样地的枯落物层渗漏水中,其TOC浓度彼此相差不大,平均为24.51 mg/L,高于林冠穿透水的TOC浓度;受枯落物截持部分降水及与枯落物TOC交换的影响,4个样地枯落物渗漏水的TOC通量(kg/hm2)(桦木次生林84.35、野李子灌丛129.35、辽东栎次生林79.21、油松人工林114.93)都比其林下降水TOC通量增加了,但华北落叶松人工林和华山松次生林的TOC通量分别降至90.76和104.90 kg/hm2。在测定的华北落叶松人工林和华山松次生林的主根系层(0—30 cm)土壤渗漏水中,TOC浓度均低于枯落物渗漏水;由于水量减小和与土壤发生碳交换,土壤渗漏水的TOC通量均显著低于枯落物渗漏水,两个林分样地分别降至43.04和66.33 kg/hm2。整体来看,林外降水携带的TOC输入通量在林地TOC输入中占有重要地位,林冠的TOC淋洗使其程度不同地增加TOC通量,枯落物层具有增加或减少TOC通量的作用,但主根系层土壤会显著减少TOC输出通量,所以是固定TOC的重要场所。