枯落物处理对格木林土壤碳氮转化和微生物群落结构的短期影响

以我国南亚热带格木人工纯林为研究对象,采用气压过程分离(BaPS)技术和磷脂脂肪酸(PLFAs)法研究了不同枯落物处理(对照、枯落物去除、枯落物加倍)下土壤碳氮转化速率和微生物群落结构的季节变化.结果表明:不同枯落物处理土壤呼吸和总硝化速率均呈现明显的季节动态,雨季显著高于旱季.枯落物处理初期,土壤呼吸和总硝化速率均随枯落物输入量的增加呈下降趋势,但随着枯落物处理时间的延长,二者随枯落物输入量的增加而增加.旱季不同枯落物处理土壤微生物PLFAs总量和各菌群PLFAs量均显著高于雨季,而雨季真菌PLFAs/细菌PLFAs明显高于旱季.在旱季,枯落物去除处理土壤微生物PLFAs总量、细菌PLFAs量、真菌PLFAs量和丛枝菌根真菌PLFAs量分别显著提高30.9%、28.8%、44.4%和31.6%.在雨季,枯落物去除处理细菌PLFAs量和丛枝菌根真菌PLFAs量分别显著降低10.6%和33.3%.土壤微生物群落结构受枯落物输入量处理和季节的双重影响,土壤微生物群落结构主要受土壤温度和铵态氮的影响.枯落物输入量处理在短期内显著影响了格木林土壤碳氮转化速率和微生物群落结构,这种影响因季节的不同而存在差异.     

不同坡向凋落物分解对土壤微生物群落的影响

采用空间与时间序列并用的方法,对贵州茂兰喀斯特森林阳坡和阴坡凋落物的分解特征及土壤微生物进行1年的野外调查和测定,探讨凋落物分解规律及其对土壤微生物群落的影响。凋落物在分解1年后的质量损失率约为72%,在分解过程中的碳、磷元素表现为释放,氮元素表现出富集;土壤微生物各群落磷脂脂肪酸(PLFA)在分解过程中的平均含量表现为:细菌(149.8 nmol/g)放线菌(63.9 nmol/g)真菌(31.3 nmol/g),阳坡和阴坡的微生物总PLFA在180d(559.2 nmol/g)和360d(513.6 nmol/g)的含量显著高于分解前;随分解时间的增加,阳坡细菌特征磷脂脂肪酸18:1ω7c的含量增加最大,其次是cy19:0和真菌特征磷脂脂肪酸18:1ω9c,阴坡土壤微生物PLFA含量表现出减少的趋势,细菌特征脂肪酸18:1ω7c减少量最大,约9.42 nmol/g,其次是i15:0和真菌特征脂肪酸18:1ω9c,约减少4.29—4.86 nmol/g;凋落物养分释放特征与微生物群落间的Spearman相关分析表明,真菌群落与碳元素释放率呈显著正相关关系,细菌群落与凋落物碳、磷元素的释放呈极显著正相关关系,且阳坡凋落物分解对微生物群落的影响高于阴坡。坡向和凋落物养分释放量对喀斯特森林凋落物的分解及土壤微生物群落的影响较显著,特征磷脂脂肪酸18:1ω7c、cy19:0、i15:0和18:1ω9c对环境变化的响应较为敏感。     

中亚热带地区两种森林植被类型土壤微生物群落结构

利用磷脂脂肪酸(PLFA)生物标记法分析了中亚热带地区罗浮栲天然林和相邻的杉木人工林土壤微生物群落结构特点.结果表明: 两种植被类型的磷脂脂肪酸总量、细菌特征脂肪酸、真菌特征脂肪酸、放线菌特征脂肪酸、革兰氏阳性菌和革兰氏阴性菌特征脂肪酸含量均为0～10 cm高于10～20 cm土层,罗浮栲天然林高于杉木人工林.在两种植被类型的两个土层中,细菌PLFAs含量均显著高于真菌PLFAs含量.两种植被类型中,细菌PLFAs含量约占PLFAs总量的44%～52%,而真菌仅占6%～8%,表明细菌在该地区两种植被类型土壤中处于优势地位.主成分分析表明,土壤微生物群落结构差异主要由植被类型差异引起,土层深度的影响相对较小.相关分析显示,革兰氏阴性菌、革兰氏阳性菌以及细菌的PLFAs含量与pH呈显著负相关,与含水量呈显著正相关;土壤微生物主要类群PLFAs含量与总氮、有机碳、C/N和铵态氮均呈显著正相关.     

培养条件下枯落物分解过程中微生物残体对土壤有机碳形成的贡献

采用黄土丘陵区多年生C3草本植物长芒草为对象,模拟“枯落物-土壤”转换界面,进行了为期512 d的室内分解试验,对枯落物分解过程中界面土层微生物残体和土壤碳组分动态进行了研究。结果表明：土壤微生物残体的形成在分解早期和中期由真菌主导,而在晚期由细菌主导。真菌残体碳对矿物结合态有机碳的贡献率(38.7%～75.8%)明显高于细菌(9.2%～22.5%),是细菌残体贡献率的3～4倍。土壤有机碳含量在枯落物分解过程中呈下降趋势。植物碳源的输入调动了微生物对土壤碳组分的利用。颗粒态有机碳分解早期和晚期持续下降,成为土壤有机碳含量减少的直接原因;而微生物残体碳和矿质结合态有机碳的波动变化对土壤有机碳含量的降低只起到间接作用。一次性外源添加枯落物引起的土壤微生物残体碳的增加并没有直接贡献土壤有机碳的积累。     

外来种湿地松凋落物对土壤微生物群落结构和功能的影响

外来种湿地松是我国亚热带地区大面积造林树种,研究其凋落物影响下土壤微生物群落结构和功能的变化,对于全面科学的评价湿地松的生态影响具有重要的理论与实践意义。通过凋落物袋 小盆模拟试验研究,分两个时期(分解5个月和18个月)比较了外来种湿地松与本地种马尾松的凋落物对土壤微生物群落结构(磷脂脂肪酸)和功能(碳代谢)的影响,结果表明：(1)外来种湿地松凋落物的C:N高于本地种马尾松;(2)两个时期,湿地松凋落物处理土壤细菌和放线菌的磷脂脂肪酸含量均低于马尾松,18个月时湿地松凋落物处理土壤真菌含量和群落真菌/细菌显著高于马尾松处理;(3)湿地松凋落物影响下土壤微生物群落功能多样性显著低于马尾松;(4)土壤微生物群落的结构显著影响微生物的活性和功能多样性：土壤微生物群落碳源代谢的强度、多样性及丰富度与细菌磷脂脂肪酸含量呈极显著正相关,细菌特征脂肪酸14:0、15:0、a15:0、i16:0、16:1ω7c、a17:0和cy19:0的含量显著影响土壤微生物群落碳代谢功能。上述结果表明：与本地种马尾松相比,引进种湿地松的凋落物显著改变了土壤微生物群落结构,降低了微生物群落的功能。     

种植阔叶树种和毛竹对土壤有机碳矿化与微生物群落结构的影响

为探明种植阔叶树种和毛竹对土壤有机碳矿化与微生物群落特征的影响,本研究通过盆栽试验和室内培养法比较分析种植香樟、木荷、青冈等阔叶树种与毛竹的土壤有机碳矿化速率和累计矿化量,并结合末端限制性片段长度多态性(T-RFLP)以及荧光定量PCR技术,分析土壤细菌、真菌群落组分与数量特征。结果表明: 与种植阔叶树种的土壤相比,种植毛竹的土壤活性碳组分和碳矿化总量显著提高,且其地下生物量高于阔叶树种。种植毛竹的土壤真菌/细菌丰度比也高于种植阔叶树种的土壤,土壤细菌群落组分对树种的响应比真菌群落更加敏感。种植毛竹的土壤真菌群落多样性显著低于阔叶树种;与细菌群落相比,真菌群落多样性与pH、有机碳含量、碳矿化速率更为相关。与阔叶树种相比,种植毛竹能显著增加土壤碳矿化总量,而毛竹土壤真菌群落结构是导致碳矿化量较高的原因之一。     

庞泉沟自然保护区常见禾本科植物内生真菌分布及其影响因素

以庞泉沟自然保护区4种不同群落类型林地和灌丛下的优势禾草为研究对象,调查不同禾草内生真菌感染率,研究内生真菌感染的禾草种群与原生生境的土壤理化性质、土壤酶活性及土壤微生物群落结构的相互关系。结果发现,影响紫羊茅、中亚薹草和野燕麦染菌率的主要因子是土壤水分含量和碳氮比,且土壤C、N、S以及土壤水分含量与紫羊茅的内生真菌染菌率成正相关,土壤蔗糖酶、脲酶和酸性磷酸酶与紫羊茅的感染率显著正相关,但中亚薹草与紫羊茅的结果相反。丝盖伞科是华北落叶松林和油松林土壤微生物群落的优势真菌,但不同染菌植被群落类型下的优势细菌微生物群落组成不同。总体看来,4种森林群落类型下禾草内生真菌染菌率与土壤理化性质相关性不同,且受到的生态因子影响大小也有差异,内生真菌感染导致各植被群落类型的土壤微生物群落结构具有差异,其优势土壤真菌和土壤细菌组成不同,这有助于进一步认识原生生境下不同感染率的禾草种群对土壤微生物群落结构的影响及其在生态系统中的各异生态学功能。     

中亚热带地区米槠天然林土壤微生物群落结构的多样性

为了解土壤微生物群落的结构,采用磷脂脂肪酸方法对武夷山和建瓯的米槠(Castanopsis carlesii)天然林土壤微生物群落的结构多样性进行了研究。结果表明,两地米槠天然林的土壤微生物群落组成十分丰富,多样性指数、丰富度指数和均匀度指数分别为2.92~3.01、25.84~28.23 和0.88~0.90。0~10 cm土层的磷脂脂肪酸总量、细菌特征脂肪酸、真菌特征脂肪酸、放线菌特征脂肪酸、革兰氏阳性菌和阴性菌特征脂肪酸含量均高于10~20 cm土层的,且建瓯万木林自然保护区的高于武夷山国家级自然保护区。10~20 cm土层的革兰氏阳性菌/革兰氏阴性菌高于0~10 cm土层的;细菌特征脂肪酸含量显著高于真菌,表明细菌在土壤微生物群落结构中处于优势地位。主成分分析表明,土壤微生物群落结构的差异主要是由采样地点的不同引起。     

模拟根系分泌物C:N化学计量特征对川西亚高山森林土壤碳动态和微生物群落结构的影响

目前有关森林根系分泌物及其诱导的土壤生态学效应研究主要关注根系碳(C)源输入, 而极少关注根系分泌物氮(N)源输入及其伴随的C:N化学计量特征对土壤过程和功能的影响, 极大地限制了我们对森林根系-土壤-微生物互作机制的深入认识。该研究以川西亚高山天然林和云杉(Picea asperata)人工林土壤为对象, 模拟配制不同C:N化学计量特征(只有N、C:N = 10、C:N = 50、C:N = 100和只有C处理)的根系分泌物溶液进行人工添加试验, 以探究根系分泌物化学计量特征对两种林分土壤碳动态及其微生物群落结构的影响差异。结果表明: 模拟根系分泌物C添加总体促进了两种林分土壤有机质分解激发效应而降低了土壤总碳(TC)含量, 而N添加在一定程度上缓和了两种林分土壤TC含量的降低幅度, 且C添加导致天然林土壤TC含量的降低幅度明显低于土壤N有效性更低的人工林。几种根系分泌物添加处理对两种林分土壤活性和惰性碳库的影响无明显规律。另外, 根系分泌物C添加总体降低了天然林土壤微生物总磷脂脂肪酸(PLFA)含量和细菌、放线菌、真菌PLFA含量, 而总体增加人工林土壤微生物PLFA总量和细菌、放线菌、真菌PLFA含量, 并诱导两种林分土壤微生物群落结构(细菌:真菌相对丰度)也发生了各自不同的变化。上述结果表明森林根系分泌物N源输入和土壤N有效性共同调控根系C源输入对土壤有机质分解激发效应的方向和幅度。研究结果为深入揭示典型森林根系分泌物化学计量特征对土壤生物化学循环过程的调控机制提供了一定的理论依据。     

Effects of simulated exudate C:N stoichiometry on dynamics of carbon and microbial community composition in a subalpine coniferous forest of western Sichuan,China

 目前有关森林根系分泌物及其诱导的土壤生态学效应研究主要关注根系碳(C)源输入, 而极少关注根系分泌物氮(N)源输入及其伴随的C:N化学计量特征对土壤过程和功能的影响, 极大地限制了我们对森林根系-土壤-微生物互作机制的深入认识。该研究以川西亚高山天然林和云杉(Picea asperata)人工林土壤为对象, 模拟配制不同C:N化学计量特征(只有N、C:N = 10、C:N = 50、C:N = 100和只有C处理)的根系分泌物溶液进行人工添加试验, 以探究根系分泌物化学计量特征对两种林分土壤碳动态及其微生物群落结构的影响差异。结果表明: 模拟根系分泌物C添加总体促进了两种林分土壤有机质分解激发效应而降低了土壤总碳(TC)含量, 而N添加在一定程度上缓和了两种林分土壤TC含量的降低幅度, 且C添加导致天然林土壤TC含量的降低幅度明显低于土壤N有效性更低的人工林。几种根系分泌物添加处理对两种林分土壤活性和惰性碳库的影响无明显规律。另外, 根系分泌物C添加总体降低了天然林土壤微生物总磷脂脂肪酸(PLFA)含量和细菌、放线菌、真菌PLFA含量, 而总体增加人工林土壤微生物PLFA总量和细菌、放线菌、真菌PLFA含量, 并诱导两种林分土壤微生物群落结构(细菌:真菌相对丰度)也发生了各自不同的变化。上述结果表明森林根系分泌物N源输入和土壤N有效性共同调控根系C源输入对土壤有机质分解激发效应的方向和幅度。研究结果为深入揭示典型森林根系分泌物化学计量特征对土壤生物化学循环过程的调控机制提供了一定的理论依据。     

Connecting plant–microbial interactions above and belowground: a fungal endophyte affects decomposition

Mutualisms can strongly affect the structure of communities, but their influence on ecosystem processes is not well resolved. Here we show that a plant–microbial mutualism affects the rate of leaf litter decomposition using the widespread interaction between tall fescue grass (Lolium arundinaceum) and the fungal endophyte Neotyphodium coenophialum. In grasses, fungal endophytes live symbiotically in the aboveground tissues, where the fungi gain protection and nutrients from their host and often protect host plants from biotic and abiotic stress. In a field experiment, decomposition rate depended on a complex interaction between the litter source (collected from endophyte-infected or endophyte-free plots), the decomposition microenvironment (endophyte-infected or endophyte-free plots), and the presence of mesoinvertebrates (manipulated by the mesh size of litter bags). Over all treatments, decomposition was slower for endophyte-infected fescue litter than for endophyte-free litter. When mesoinvertebrates were excluded using fine mesh and litter was placed in a microenvironment with the endophyte, the difference between endophyte-infected and endophyte-free litter was strongest. In the presence of mesoinvertebrates, endophyte-infected litter decomposed faster in microenvironments with the endophyte than in microenvironments lacking the endophyte, suggesting that plots differ in the detritivore assemblage. Indeed, the presence of the endophyte in plots shifted the composition of Collembola, with more Hypogastruridae in the presence of the endophyte and more Isotomidae in endophyte-free plots. In a separate outdoor pot experiment, we did not find strong effects of the litter source or the soil microbial/microinvertebrate community on decomposition, which may reflect differences between pot and field conditions or other differences in methodology. Our work is among the first to demonstrate an effect of plant–endophyte mutualisms on ecosystem processes under field conditions.     

内生真菌对氮添加羽茅根际土壤特性和微生物群落的影响

内生真菌不仅能改变与其共生植物的生理和生长指标,还可通过宿主植物间接对土壤的理化性质和微生物群落结构产生影响。以天然禾草——羽茅(Achnatherum sibiricum)为研究材料,探究在不同施氮水平下,内生真菌的种类对不同基因型的宿主植物根际土壤理化性质和微生物群落产生何种影响。结果表明,内生真菌侵染显著提高了羽茅根际土壤的pH值和微生物总量,但降低了土壤中真菌与细菌的比值。同时,土壤的pH值还受到了内生真菌种类的影响,其中感染Epichlo3 sibirica的羽茅根际土壤pH显著高于感染Epichlo3 gansuense-1的羽茅,而感染Epichlo3 gansuense-2的羽茅根际土壤pH与感染E.gansuensis-1、E.sibirica菌的羽茅相比没有显著差异。另外,内生真菌感染与否、内生真菌种类、施氮量以及宿主植物基因型对土壤总碳、总氮、微生物及碳矿化能力均无显著影响。     

How does the Fungal Endophyte Neotyphodium coenophialum Affect Tall Fescue (Festuca arundinacea) Rhizodeposition and Soil Microorganisms?

The goal of our study was to investigate the impact of fungal endophytes in tall fescue (Festuca arundinacea) on rhizodeposition and in turn, the soil microbial community. Sand-based, aseptic microlysimeter units were constructed for the collection of rhizodeposit solutions for chemical analyses from the roots of endophyte-free (E−) and endophyte-infected (E+) tall fescue plants. E+ plants were infected with Neotyphodium coenophialum, the most common endophyte found in tall fescue. Rhizodeposit solutions collected over nine weeks from E+ grass contained more organic carbon and carbohydrates than E−. These solutions were allowed to percolate through columns of plant-free soils to assess the response of the soil microbial communities. Soils to which solutions from E+ grass were applied had significantly higher respiration rates than those receiving solutions from E− grass, suggesting that microbial activity was stimulated by changes in the rhizodeposits. Culture-based assays of the soil microbial community (plate counts and community-level physiological profiling) suggest that the basic structure of the microbial community was not affected by application of rhizodeposit solutions from E+ plants as compared to E−. Our results indicate that the presence of a fungal endophyte may enhance rhizodeposition by tall fescue and could consequently influence microbial mineralization processes in the soil. In grasslands where nutrients may be limiting, hosting a fungal endophyte has the potential to enhance plant nutrient supply indirectly via a stimulatory effect on the soil microbial biomass. Megan M. Van Hecke and Amy M. Treonis - Both authors contributed equally to this work.     

Do foliar endophytes affect grass litter decomposition? A microcosm approach using Lolium multiflorum

Symbiotic infection with fungal endophytes has been shown to decrease herbivory in several temperate grasses. We tested the hypothesis that foliar endophytes of grasses may also affect below-ground processes upon their host death, by altering the litter quality for detritivores or the microenvironment for decomposition. Microcosm–litterbag experiments were used to assess decay rates for litter produced by endophyte ( Neotyphodium sp.) infected vs uninfected Lolium multiflorum plants, and to examine endophyte-mediated effects of prior site occupants on current litter decomposition. We found that litter from endophyte-infected L. multiflorum decomposed more slowly than litter from endophyte-free conspecifics and from a naturally uninfected grass, Bromus unioloides . Furthermore, the endophyte–grass association modified the decomposition environment, so that B. unioloides litter decomposed faster when placed underneath a thick layer of endophyte-free L. multiflorum litter. Litter decay rates increased with the amount of root debris remaining in situ from the previous season, but were not affected by the infection status of prior site occupants. The lower decomposability of litter from infected L. multiflorum plants persisted across a range of microenvironments, as determined by different amounts of above-ground litter and soil moisture conditions. Endophyte infection tended to reduce the N content of decaying litter; however, litter N and C/N ratio mainly accounted for interspecific differences in decomposition. Our results imply that fungal endophytes not only can affect herbivory food chains, but also soil organisms and the ecosystem processes they regulate. This study suggests a novel role for symbiotic foliar endophytes in linking above-ground and below-ground sub-systems.     

Role of foliar fungal endophytes in litter decomposition among species and population origins

Litter decomposition is a key ecosystem process which returns nutrients from dead plant material to mineral forms in the soil. We examined whether systemic fungal endophytes modulate recycling of nutrients directly by altering litter decomposition. We studied litter decomposition mediated by Epichloë endophytes in litter-bag experiments. We examined direct endophyte effects on litter decomposition in wild populations and cultivars of Schedonorus phoenix and Schedonorus pratensis. In the first experiment, endophyte presence tended to increase litter decomposition rate in cultivars of the two grass species (S. phoenix and S. pratensis). However, in the second experiment plant origin had a stronger influence than endophyte symbiosis in S. phoenix. Interestingly, the initial level of alkaloids was associated positively with decomposition in S. phoenix populations. Characteristics associated with litter quality were not clearly related to either endophytes or decomposition rate. Our results suggest that endophytes can enhance litter breakdown but their role in nutrient cycling is far more complex depending on plant population origin.     

Interactions between an arbuscular mycorrhizal fungus and a soil microbial community mediating litter decomposition

We investigated arbuscular mycorrhizal fungi (AMF) alteration of microbial mediation of litter decomposition. AMF (Glomus hoi) were either allowed access to or excluded from Plantago lanceolata L. root litter embedded in soil; litter was labeled with either (13) C only or (13) C and (15) N. Plant N uptake was significantly increased if AMF accessed the litter, and (15) N analysis of the plant material indicated that 2-3% of plant N originated from litter. Succession of the soil community mediating decomposition was assessed by phospholipid fatty acids (PLFA) combined with (13) C-PLFA. During the first 21?days of decomposition, saprotrophic fungi and Gram-negative bacteria were the dominant consumers of litter C. As decomposition progressed however, (13) C content of the fungal biomarkers declined substantially, and Gram-negative and Gram-positive bacteria became the primary reservoirs of labeled litter C. The putative PLFA marker for AMF (16:1ω5c) originated primarily from non-AMF sources. In AMF-invaded root litter, Gram-negative, Gram-positive, and 16:1ω5c markers became less (13) C-enriched relative to their counterparts in non-AMF-invaded microcosms during active decomposition. These patterns of (13) C: (12) C enrichment may result from AMF supply of (12) C from the plant to the decomposing soil microbial community; such C inputs could alter the microbial mediation of litter decomposition.     

Direct and indirect effects of the fungal endophyte <Emphasis Type="Italic">Epichloë uncinatum</Emphasis> on litter decomposition of the host grass, <Emphasis Type="Italic">Schedonorus pratensis</Emphasis>

Microbial plant symbionts have been suggested to mediate plant-soil feedback and affect ecosystem functions. Systemic Epichloë fungal endophytes of grasses are found to mediate litter decomposition. These effects are often linked to alkaloids produced by Epichloë species, which are hypothesized to negatively affect decomposers. Although endophytes have been found to affect plant community and soil biota, direct (through litter quality) and indirect (through the environment) effects of fungal endophytes on litter decomposition have been scarcely scrutinized. We placed litterbags with endophyte-symbiotic (E+) and non-symbiotic (E?) Schedonorus pratensis plant litter in plots dominated by E+ or E? plants of the same species, and followed the dynamics of mass losses over time. We predicted the endophyte would hinder decomposition through changes in litter quality and that both types of litter would decompose faster in home environments. E+ litter decomposed faster in both environments. The mean difference between decomposition rate of E+ and E? litter tended to be higher in E? plots. Nitrogen and phosphorus, two elements usually associated with high decomposition rates, were significantly lower in E+ litter. We also detected a higher proportion of C in the cellulose form in E+ litter. Contrary to the general assumption, we found that symbiosis with Epichloë fungal endophytes can be associated with higher decomposition of plant litter. Since direct effects of Epichloë fungi were still stronger than indirect effects, it is suggested that besides the alkaloids, other changes in plant biomass would explain in a context-dependent manner, the endophyte effects on the litter decomposition.     

氮肥添加对高寒藏嵩草(Kobresia tibetica)沼泽化草甸和土壤微生物群落的影响

以藏嵩草沼泽化草甸为研究对象,利用磷脂脂肪酸(PLFA)技术,研究连续6年N素添加对地上植被群落数量特征、土壤微生物群落结构的影响。结果表明:①藏嵩草沼泽化草甸群落生物量、枯枝落叶对施肥处理无明显响应,且莎草科植物对土壤氮素的吸收和利用率较低。②施肥增加了0-10 cm土壤微生物类群PLFAs丰富度尤其细菌和革兰氏阳性菌PLFAs,降低了10-20 cm PLFAs丰富度;③磷脂脂肪酸饱和脂肪酸/单烯不饱和脂肪酸、细菌PLFAs/真菌PLFAs的比值随土壤层次增加而增加;④0-10 cm土层革兰氏阳性菌、真菌PLFAs含量与pH、土壤速效磷、速效氮、土壤有机质显著正相关(P0.05或P0.01);10-20 cm土层,细菌、革兰氏阳性菌、真菌和总PLFAs含量与土壤有机质含量显著正相关(P0.05或P0.01)。表明藏嵩草沼泽化草甸微生物PLFAs含量和丰富度对施肥的响应存在明显的土层梯度效应,土壤微生物PLFAs含量和丰富度主要受表层土壤初始养分含量的影响。