Salinity controls soil microbial community structure and function in coastal estuarine wetlands

Soil salinity acts as a critical environmental filter on microbial communities, but the consequences for microbial diversity and biogeochemical processes are poorly understood. Here, we characterized soil bacterial communities and microbial functional genes in a coastal estuarine wetland ecosystem across a gradient (~5 km) ranging from oligohaline to hypersaline habitats by applying the PCR-amplified 16S rRNA (rRNA) genes sequencing and microarray-based GeoChip 5.0 respectively. Results showed that saline soils in marine intertidal and supratidal zone exhibited higher bacterial richness and Faith's phylogenetic diversity than that in the freshwater-affected habitats. The relative abundance of taxa assigned to Gammaproteobacteria, Bacteroidetes and Firmicutes was higher with increasing salinity, while those affiliated with Acidobacteria, Chloroflexi and Cyanobacteria were more prevalent in wetland soils with low salinity. The phylogenetic inferences demonstrated the deterministic role of salinity filtering on the bacterial community assembly processes. The abundance of most functional genes involved in carbon degradation and nitrogen cycling correlated negatively with salinity, except for the hzo gene, suggesting a critical role of the anammox process in tidal affected zones. Overall, the salinity filtering effect shapes the soil bacterial community composition, and soil salinity act as a critical inhibitor in the soil biogeochemical processes in estuary ecosystems.     

Damming river shapes distinct patterns and processes of planktonic bacterial and microeukaryotic communities

Planktonic bacterial and microeukaryotic communities play important roles in biogeochemical cycles, but their biogeographic patterns and community assembly processes in large damming rivers still remain unclear. In this study, 16S rRNA and 18S rRNA coding genes were used for sample sequencing analysis of planktonic bacterial and microeukaryotic communities in the upper Yangtze River. The upper Yangtze River was divided into dam-affected zones and river zones based on the influence of dams. The results showed that there were significant differences in the bacterial and microeukaryotic communities between the two zones and that dams significantly reduced the α-diversity of the bacterial communities. Co-occurrence network analysis indicated that networks in the river zone were denser than those in the dam-affected zone. The relationships among species in bacterial networks were more complex than those in microeukaryotic networks. Dispersal limitation and ecological drift were the main processes influencing planktonic bacterial and microeukaryotic communities in the dam-affected zone respectively, whereas the role of deterministic processes increased in the river zone. Anthropogenic activities and hydraulic conditions affected suspended sediment and controlled microbial diversity in the river zone. These results suggest that dams impact planktonic bacteria more strongly than planktonic microeukaryotes, indicating that the distribution patterns and processes of the bacterial and microeukaryotic communities in large rivers are significantly different.     

猫儿山不同海拔植被带土壤微生物群落功能多样性

为研究中亚热带森林土壤微生物群落功能多样性特征及其随海拔梯度的变化,应用Biolog微平板技术,对猫儿山不同海拔植被带(常绿阔叶林(EBF)、落叶阔叶混交林(DBF)、针阔混交林(CBF))土壤微生物群落功能多样性差异进行了比较。结果表明,不同海拔植被带土壤微生物群落功能多样性差异显著。土壤平均颜色变化率(AWCD)随培养时间延长而逐渐增加,随着海拔升高,土壤AWCD值逐渐降低,大小顺序为EBFDBFCBF。土壤微生物群落Shannon指数和丰富度指数的总体趋势为EBF最高,DBF次之,CBF最低。不同海拔植被带土壤微生物群落均匀度指数之间差异不显著。不同海拔植被带土壤微生物对不同碳源的利用能力存在差异,其中EBF利用率最高,CBF利用率最低,氨基酸类、胺类和酯类碳源为各海拔植被带土壤微生物利用的主要碳源。主成分分析结果表明,主成分1和主成分2分别能解释变量方差的40.42%和15.97%,在主成分分离中起主要贡献作用的是酯类、胺类和氨基酸类碳源。土壤理化性质与土壤微生物群落功能多样性之间的相关性分析结果表明,微生物群落多样性的Shannon指数与全钾(TK)呈极显著正相关(P0.01),与含水量呈极显著负相关(P0.01),与总有机碳(TOC)、全氮(TN)、速效氮(AN)、有效P(AP)之间的相关性显著(P0.05)或极显著(P0.01),且为负相关。土壤TK含量和含水量可能是造成不同海拔土壤微生物群落功能多样性差异的主要原因。     

Shifts of tundra bacterial and archaeal communities along a permafrost thaw gradient in Alaska

Understanding the response of permafrost microbial communities to climate warming is crucial for evaluating ecosystem feedbacks to global change. This study investigated soil bacterial and archaeal communities by Illumina MiSeq sequencing of 16S rRNA gene amplicons across a permafrost thaw gradient at different depths in Alaska with thaw progression for over three decades. Over 4.6 million passing 16S rRNA gene sequences were obtained from a total of 97 samples, corresponding to 61 known classes and 470 genera. Soil depth and the associated soil physical–chemical properties had predominant impacts on the diversity and composition of the microbial communities. Both richness and evenness of the microbial communities decreased with soil depth. Acidobacteria, Verrucomicrobia, Alpha‐ and Gamma‐Proteobacteria dominated the microbial communities in the upper horizon, whereas abundances of Bacteroidetes, Delta‐Proteobacteria and Firmicutes increased towards deeper soils. Effects of thaw progression were absent in microbial communities in the near‐surface organic soil, probably due to greater temperature variation. Thaw progression decreased the abundances of the majority of the associated taxa in the lower organic soil, but increased the abundances of those in the mineral soil, including groups potentially involved in recalcitrant C degradation (Actinomycetales, Chitinophaga, etc.). The changes in microbial communities may be related to altered soil C sources by thaw progression. Collectively, this study revealed different impacts of thaw in the organic and mineral horizons and suggests the importance of studying both the upper and deeper soils while evaluating microbial responses to permafrost thaw.     

The balance between deterministic and stochastic processes in structuring lake bacterioplankton community over time

One major goal in microbial ecology is to establish the importance of deterministic and stochastic processes for community assembly. This is relevant to explain and predict how diversity changes at different temporal scales. However, understanding of the relative quantitative contribution of these processes and particularly of how they may change over time is limited. Here, we assessed the importance of deterministic and stochastic processes based on the analysis of the bacterial microbiome in one alpine oligotrophic and in one subalpine mesotrophic lake, which were sampled over two consecutive years at different time scales. We found that in both lakes, homogeneous selection (i.e., a deterministic process) was the main assembly process at the annual scale and explained 66.7% of the bacterial community turnover, despite differences in diversity and temporal variability patterns between ecosystems. However, in the alpine lake, homogenizing dispersal (i.e., a stochastic process) was the most important assembly process at the short‐term (daily and weekly) sampling scale and explained 55% of the community turnover. Alpha diversity differed between lakes, and seasonal stability of the bacterial community was more evident in the oligotrophic lake than in the mesotrophic one. Our results demonstrate how important forces that govern temporal changes in bacterial communities act at different time scales. Overall, our study validates on a quantitative basis, the importance and dominance of deterministic processes in structuring bacterial communities in freshwater environments over long time scales.     

Niche partitioning in arbuscular mycorrhizal communities in temperate grasslands: a lesson from adjacent serpentine and nonserpentine habitats

Arbuscular mycorrhizal fungi (AMF) represent an important soil microbial group playing a fundamental role in many terrestrial ecosystems. We explored the effects of deterministic (soil characteristics, host plant life stage, neighbouring plant communities) and stochastic processes on AMF colonization, richness and community composition in roots of Knautia arvensis (Dipsacaceae) plants from three serpentine grasslands and adjacent nonserpentine sites. Methodically, the study was based on 454‐sequencing of the ITS region of rDNA. In total, we detected 81 molecular taxonomical operational units (MOTUs) belonging to the Glomeromycota. Serpentine character of the site negatively influenced AMF root colonization, similarly as higher Fe concentration. AMF MOTUs richness linearly increased along a pH gradient from 3.5 to 5.8. Contrary, K and Cr soil concentration had a negative influence on AMF MOTUs richness. We also detected a strong relation between neighbouring plant community composition and AMF MOTUs richness. Although spatial distance between the sampled sites (c. 0.3–3 km) contributed to structuring AMF communities in K. arvensis roots, environmental parameters were key factors in this respect. In particular, the composition of AMF communities was shaped by the complex of serpentine conditions, pH and available soil Ni concentration. The composition of AMF communities was also dependent on host plant life stage (vegetative vs. generative). Our study supports the dominance of deterministic factors in structuring AMF communities in heterogeneous environment composed of an edaphic mosaic of serpentine and nonserpentine soils.     

Changes in assembly processes in soil bacterial communities following a wildfire disturbance

Although recent work has shown that both deterministic and stochastic processes are important in structuring microbial communities, the factors that affect the relative contributions of niche and neutral processes are poorly understood. The macrobiological literature indicates that ecological disturbances can influence assembly processes. Thus, we sampled bacterial communities at 4 and 16 weeks following a wildfire and used null deviation analysis to examine the role that time since disturbance has in community assembly. Fire dramatically altered bacterial community structure and diversity as well as soil chemistry for both time-points. Community structure shifted between 4 and 16 weeks for both burned and unburned communities. Community assembly in burned sites 4 weeks after fire was significantly more stochastic than in unburned sites. After 16 weeks, however, burned communities were significantly less stochastic than unburned communities. Thus, we propose a three-phase model featuring shifts in the relative importance of niche and neutral processes as a function of time since disturbance. Because neutral processes are characterized by a decoupling between environmental parameters and community structure, we hypothesize that a better understanding of community assembly may be important in determining where and when detailed studies of community composition are valuable for predicting ecosystem function.     

内蒙古贝加尔针茅草原不同利用方式土壤微生物功能多样性

基于Biolog-ECO技术,研究了贝加尔针茅草原在自由放牧、刈割和围封3种不同利用方式下土壤微生物群落功能多样性变化。结果表明:不同的利用方式能显著改变土壤微生物群落代谢活性,反映微生物活性的平均颜色变化率表现为围封>自由放牧>刈割,围封时土壤微生物群落代谢活性最高;不同的利用方式改变了土壤微生物群落多样性指数,自由放牧土壤微生物群落丰富度指数、均匀度指数和优势度指数均最高,围封次之,刈割最低。主成分分析结果表明:在自由放牧和刈割2种利用方式下土壤微生物群落碳源利用模式及代谢功能相似,而围封土壤微生物群落具有不同的碳源利用模式和代谢功能;糖类、氨基酸类和代谢中间产物及次生代谢物为土壤微生物利用的主要碳源。不同的利用方式改变了贝加尔针茅草原土壤微生物群落功能多样性。     

Crop rotational diversity enhances belowground communities and functions in an agroecosystem

Biodiversity loss, an important consequence of agricultural intensification, can lead to reductions in agroecosystem functions and services. Increasing crop diversity through rotation may alleviate these negative consequences by restoring positive aboveground–belowground interactions. Positive impacts of aboveground biodiversity on belowground communities and processes have primarily been observed in natural systems. Here, we test for the effects of increased diversity in an agroecosystem, where plant diversity is increased over time through crop rotation. As crop diversity increased from one to five species, distinct soil microbial communities were related to increases in soil aggregation, organic carbon, total nitrogen, microbial activity and decreases in the carbon‐to‐nitrogen acquiring enzyme activity ratio. This study indicates positive biodiversity–function relationships in agroecosystems, driven by interactions between rotational and microbial diversity. By increasing the quantity, quality and chemical diversity of residues, high diversity rotations can sustain soil biological communities, with positive effects on soil organic matter and soil fertility.     

土壤微生物群落对麻栎-刺槐混交林凋落物分解的影响

以麻栎-刺槐混交林和麻栎纯林为研究对象,采用野外定点采样、室内分析与高通量测序的方法,对凋落物分解过程中土壤微生物菌群多样性特征及其对凋落物分解速率的影响进行了研究。结果表明:(1)麻栎-刺槐混交林凋落物的分解速率高于麻栎纯林。两种林分凋落物有机碳(TOC)、全氮(TN)发生释放,全磷(TP)发生积累-释放的过程。(2)两种林分土壤细菌优势类群为放线菌门(Acidobacteria)、变形菌门(Proteobacteria)、酸杆菌门(Actinobacteria)和疣微菌门(Verrucomicrobia),土壤真菌优势类群为担子菌门(Basidiomycota)、子囊菌门(Ascomycota)和被孢霉门(Moritierellomycota)。(3)凋落物分解过程中,麻栎-刺槐混交林土壤微生物菌群丰富度指数和菌群多样性指数变化范围小于麻栎纯林。(4)凋落物分解速率与土壤细菌菌群丰富度指数和菌群多样性指数呈显著正相关,与土壤真菌菌群丰富度指数呈显著正相关。土壤微生物群落对麻栎-刺槐混交林和麻栎纯林凋落物分解速率具有重要影响,研究结果为深入开展混交林土壤微生物多样性对凋落物分解的影响研究提供理论依据。     

包头市半干旱型森林公园土壤细菌多样性与功能

土壤微生物是城市公园生态系统中物质循环的主要驱动者, 但目前对于半干旱型城市公园森林土壤细菌群落结构和功能研究较少。本研究以包头市为例, 选取奥林匹克公园(AL)、劳动公园(LD)和阿尔丁植物园(ZW) 3个典型的森林公园, 利用Illumina高通量测序的方法对细菌16S rRNA V4-V5片段进行测序, 分析城市森林公园土壤细菌的多样性、群落构成以及背后的生态学构建机制, 并使用Tax4Fun对细菌群落代谢功能进行分析。结果表明, 土壤细菌丰富度指数为LD (2,443.00 ± 9.37) > ZW (2,392.90 ± 8.23) > AL (2,305.57 ± 17.48); 细菌群落以放线菌门、变形菌门、酸杆菌门、绿弯菌门和芽单胞菌门为优势门。细菌群落组成在三个公园间存在显著差异, 同时线性判别分析效应大小(LEfSe)结果表明, 所有公园都具有多度显著差异的细菌可操作分类单元(OTUs); 中性群落模型(NCM)、标准化随机率(NST)和基于系统发育结合的零模型推断群落组装机制(iCAMP)分析显示, 城市森林土壤细菌群落的构建是由随机性过程和确定性过程共同驱动, 其中漂变和同质化选择占主导作用。Tax4Fun功能预测结果表明涉及膜运输、碳水化合物代谢、氨基酸代谢、能量代谢和信号转导的代谢是该城市公园林下土壤微生物的主要代谢功能, 进一步分析表明微生物群落的代谢功能在不同公园之间存在很大差异。本研究初步探究了半干旱型城市公园细菌的多样性、群落构建机制及其功能, 为城市公园绿地建设以及生态改善等提供一定的科学依据和实践指导。     

Predators temper the relative importance of stochastic processes in the assembly of prey metacommunities

Communities assemble through a combination of stochastic processes, which can make environmentally similar communities divergent (high β-diversity), and deterministic processes, which can make environmentally similar communities convergent (low β-diversity). Top predators can influence both stochasticity (e.g. colonization and extinction events) and determinism (e.g. size of the realized species pool), in community assembly, and thus their net effect is unknown. We investigated how predatory fish influenced the scaling of prey diversity in ponds at local and regional spatial scales. While fish reduced both local and regional richness, their effects were markedly more intense at the regional scale. Underlying this result was that the presence of fish made localities within metacommunities more similar in their community composition (lower β-diversity), suggesting that fish enhance the deterministic, relative to the stochastic, components of community assembly. Thus, the presence of predators can alter fundamental mechanisms of community assembly and the scaling of diversity within metacommunities.     

盐度对滨海土壤细菌多样性和群落构建过程的影响

滨海盐土是重要的农业土地后备资源。微生物是土壤中物质循环的关键动力,然而盐度对土壤微生物群落特征影响的研究还很缺乏。本研究采集滨海地区的土壤样品,研究非盐、轻盐和高盐3组不同盐度对土壤细菌数量、多样性和群落构建的影响。结果表明: 与非盐和轻盐土壤相比,高盐土壤的脱氢酶活性和细菌数量显著降低,而细菌α多样性没有变化,细菌群落结构发生分异。利用零模型反演群落构建过程,发现盐度是细菌群落构建过程的主控因子,盐度主导的高确定性过程控制了滨海盐土细菌的群落结构。说明在现有的盐度范围内,高盐土壤中同样含有丰富的微生物种质资源,具有盐土改良的生物学基础,然而由于高确定性的群落构建机制,外源物种很难定殖于滨海盐土。因此,在利用微生物技术改良滨海盐土时,应尽可能筛选耐盐的土著菌种,提高定殖效率。     

基于宏基因组技术分析全球不同温度带土壤微生物多样性

【背景】温度在塑造大尺度的土壤微生物群落方面发挥了重要作用,但目前针对全球不同温度带大尺度土壤微生物多样性方面的研究十分缺乏。【目的】明确不同温度带大尺度土壤微生物组成和功能的差异变化。【方法】从宏观的角度运用宏基因组技术对不同温度带土壤微生物群落的组成和功能进行分析。【结果】细菌的物种多样性随着温度带纬度的升高而增多,真菌的物种多样性在温带最多,在寒带最小且假丝酵母属(Candida)占绝对优势。3个温度带间除物种多样性存在差异外,微生物群落中物种丰度差异也较大,优势属和特殊属各有不同。其中值得注意的是,假单胞菌属(Pseudomonas)和芽孢杆菌属(Bacillus)的丰度在不同温度带间存在显著差异,且随着温度带纬度的升高而增多,而链霉菌属(Streptomyces)、地嗜皮菌属(Geodermatophilus)、红色杆菌属(Rubrobacter)和小单孢菌属(Micromonospora)的丰度随温度带纬度的升高而降低。在功能方面,发现与翻译后修饰、蛋白质周转、伴侣(posttranslational modification, protein turnover, chap...     

Soil fungal diversity and assembly along a xeric stress gradient in the central Namib Desert

The Namib Desert of south-western Africa is one of the oldest deserts in the world and possesses unique geographical, biological and climatic features. While research through the last decade has generated a comprehensive survey of the prokaryotic communities in Namib Desert soils, little is yet known about the diversity and function of edaphic fungal communities, and even less of their responses to aridity. In this study, we have characterized soil fungal community diversity across the longitudinal xeric gradient across the Namib desert (for convenience, divided into the western fog zone, the central low-rainfall zone and the eastern high-rainfall zone), using internal transcribed sequence (ITS) metabarcoding. Ascomycota, Basidiomycota and Chytridiomycota consistently dominated the Namib Desert edaphic fungal communities and a core mycobiome composed of only 15 taxa, dominated by members of the class Dothideomycetes (Ascomycota), was identified. However, fungal community structures were significantly different in the fog, low-rainfall and high-rainfall zones. Furthermore, Namib Desert gravel plain fungal community assembly was driven by both deterministic and stochastic processes; the latter dominating in the all three xeric zones. We also present data that suggest that the inland limit of fog penetration represents an ecological barrier to fungal dispersal across the Namib Desert.     

黄土丘陵沟壑区地形变化对土壤微生物群落功能多样性的影响

研究黄土高原丘陵沟壑区破碎地形对土壤微生物功能多样性的影响,对于理解复杂地形区生态过程与系统功能的空间变化具有重要意义。选择陕西省安塞县陈家洼为研究区,依据坡面地形变化选择不同坡位土壤,采用Biolog微平板培养法探究地形变化对土壤微生物群落功能多样性的影响。实验发现,土壤微生物群落培养的平均颜色变化率(AWCD)增长曲线总的呈现出坡下部坡中部坡上部的规律,且坡下部AWCD值与坡中部、坡上部间差异显著(P0.05);坡下部土壤微生物群落功能多样性显著高于坡中部和坡上部,但不同土层深度(0—10 cm、10—20 cm)间无显著性差异(P0.05);对土壤微生物群落功能多样性差异贡献较大的碳源是糖类、羧酸类和多酚化合物类碳源;土壤含水率高低是不同坡位土壤微生物群落功能多样性差异显著的主要原因;微生物群落丰富度(H)和均一度(D)与土壤全氮含量正相关,优势度(U)反之,土壤全碳、全磷和p H对土壤微生物群落结构和功能多样性差异作用不显著。     

Distinct patterns of soil bacterial and fungal community assemblages in subtropical forest ecosystems under warming

Climate change globally affects soil microbial community assembly across ecosystems. However, little is known about the impact of warming on the structure of soil microbial communities or underlying mechanisms that shape microbial community composition in subtropical forest ecosystems. To address this gap, we utilized natural variation in temperature via an altitudinal gradient to simulate ecosystem warming. After 6 years, microbial co-occurrence network complexity increased with warming, and changes in their taxonomic composition were asynchronous, likely due to contrasting community assembly processes. We found that while stochastic processes were drivers of bacterial community composition, warming led to a shift from stochastic to deterministic drivers in dry season. Structural equation modelling highlighted that soil temperature and water content positively influenced soil microbial communities during dry season and negatively during wet season. These results facilitate our understanding of the response of soil microbial communities to climate warming and may improve predictions of ecosystem function of soil microbes in subtropical forests.     

Deterministic processes have limited impacts on foliar fungal endophyte communities along a savanna-forest successional gradient

Patterns and drivers of succession provide insight into the mechanisms that govern community assembly, but remain poorly understood for microbial communities. We assess whether successional trends of trees are mirrored by foliar endophyte communities of three tree species across a deterministic woody successional gradient. Additionally, we test the relative contribution of abiotic predictors, biotic factors, and spatial distance between sites in predicting composition and richness of endophyte communities. Unlike the tree community, endophyte communities showed no consistent evidence of deterministic succession. Host identity was the most important factor structuring endophyte community composition; within hosts, spatial distance from the indigenous forest and between samples was important, while environmental predictors had small and inconsistent effects. Much variation in endophyte composition remained unexplained. In contrast, endophyte richness was well-explained by predictor variables. Host identity was most important in predicting endophyte richness, while the effect of other predictors on richness differed between host species. We conclude that deterministic succession in trees did not result in deterministic succession in endophyte communities; instead community assembly was most strongly influenced by host identity; while within hosts, neutral processes may be more important for endophyte assembly than deterministic factors.     

Effects of continuous cucumber cropping and alternative rotations under protected cultivation on soil microbial community diversity

The diversity of soil microbial communities as affected by continuous cucumber cropping and alternative rotations under protected cultivation were evaluated using community level physiological profiles (CLPP) and random amplified polymorphic DNA (RAPD) analysis. The soils were selected from six cucumber cropping systems, which cover two cropping practices (rotation and continuous cropping) and a wide spectrum for cucumber cropping history under protected cultivation. Shannon–Weaver index and multivariate analysis were performed to characterize variations in soil microbial communities. Both CLPP and RAPD techniques demonstrated that cropping systems and plastic-greenhouse cultivation could considerably affect soil microbial functional diversity and DNA sequence diversity. The open-field soil had the highest Shannon–Weaver index (3.27 for CLPP and 1.50 for RAPD), whereas the lowest value occurred in the 7-year continuous protected cultivation soil (3.27 for CLPP and 1.50 for RAPD). The results demonstrated that continuous plastic-greenhouse cultivation and management can cause the reduction in the species diversity of the biota. Higher Shannon–Weaver index and coefficients of DNA sequence similarity were found in soils under rotation than those under continuous cropping. Cluster analysis also indicated that microbial community profiles of continuous cultivation soils were different from profiles of rotation soils. The reduction in diversity of microbial communities found in continuous cultivation soils as compared with rotation soils might be due to the differences in the quantity, quality and distribution of soil organic matter. Section Editor: D. E. Crowley     

中亚热带森林土壤微生物群落多样性随海拔梯度的变化

运用Biolog EcoPlate技术, 对武夷山不同海拔植被带(常绿阔叶林(EBF)、针叶林(CF)、亚高山矮林(DF)、高山草甸(AM))土壤微生物群落多样性差异进行了研究。结果表明: 不同海拔植被带土壤微生物群落功能多样性差异显著。土壤平均颜色变化率(AWCD)随培养时间延长而逐渐增加, 同一深度土层的AWCD值随海拔升高而逐渐降低, 大小顺序依次为EFB > CF > DF > AM。同一海拔植被带, 不同深度土层的AWCD值总体趋势依次为0-10 cm > 10-25 cm > 25-40 cm。土壤微生物群落Simpson指数、Shannon-Wiener指数、丰富度指数和McIntosh指数的总体趋势为EBF最高, CF和DF次之, AM最低。不同海拔植被带土壤微生物对不同碳源利用强度存在较大差异, 其中EBF利用率最高, AM利用率最低, 碳水化合物和羧酸类碳源是各海拔植被带土壤微生物的主要碳源。主成分分析结果表明, 从31个因素中提取的与碳源利用相关的主成分1、主成分2分别能解释变量方差的75.27%和16.14%, 在主成分分离中起主要贡献作用的是胺类和氨基酸类碳源。土壤微生物群落多样性随着海拔上升、土层加深而逐渐下降的原因, 可能是生物量、林分凋落物、土壤养分、微小动物、植物根系等多种因素共同作用的结果。