支持物倾角对攀援植物栝楼形态和生长的影响

支持物倾角的变化将引起攀援植物“自遮荫”程度的改变 ,从而影响植物的生长和行为。以攀援植物栝楼 ( Trichosantheskirilowii)为材料 ,通过实验生态学的方法 ,研究了 4种支持物倾角下植株的生长和觅食行为的差异。结果表明 :( 1 )不同生长发育期栝楼植株形态对自遮荫差异的可塑性反应程度不一 ,不同角度攀援生长植株在生长前期均比生长后期有较敏感的形态可塑性反应。 ( 2 )自遮荫程度随支持物倾角的增大而增强 ,较强自遮荫环境下植株比较弱自荫环境下的植株有更大的形态可塑性。 ( 3)比主茎长、比叶面积、生物量对叶片和叶柄的配置在 4种攀援生长形式间差异均不显著。不同角度攀援植株主要通过改变分枝数量、分枝形态和分枝生物量配置以适应支持物倾角的变化 ,这说明 ,自遮荫对植株形态和生物量配置仅产生有限的影响。 ( 4 )分枝能力、分枝数量以及分枝生物量配置均在大角度攀援生长中最大 ,且与小角度攀援生长植株间有显著差异。 ( 5 )水平攀援生长植株主要通过增大主茎生物量投资以充分占有生境 ,而大角度攀援生长植株则主要通过分枝茎扩展以占据有利生境 ,不同攀援生长植株有不同的觅食行为。     

乌兰布和沙漠东北缘三种豆科绿肥植物生物量和养分含量及其对土壤肥力的影响

土壤肥力是限制乌兰布和沙漠东北缘沿黄灌区建立高产绿洲的主要因子之一。在内蒙古自治区磴口县试验地种植3种绿肥植物,比较它们改良土壤的效果。结果表明,1～2年生紫花苜蓿的总生物量干重为最高,而沙打旺则因其抗风沙能力强,地上部分生长较其它两种植物为好,由于植株中水分含量较高,其生物量的鲜重在三者中是最高的。2年生绿肥植物总生物量干重和养分含量的测定结果则显示,全氮和全磷含量的变化趋势为沙打旺〉紫花苜蓿〉草木樨,而全钾含量则为紫花苜蓿〉草木樨〉沙打旺;从3种植物对土壤改良效果来看,改良作用为沙打旺〉紫花苜蓿〉草木樨。与两种常用的绿肥作物相比,沙打旺因其对风沙较强的抗性而具有较高的改良土壤能力,因此在同类型的沙漠绿洲开发初期可以作为改良土壤的首选植物。     

人工固沙区与流沙区准噶尔无叶豆种群数量特征与空间格局对比研究

准噶尔无叶豆(Eremosparton songoricum) 是中亚荒漠特有小半灌木,稀有种,在我国仅片段化分布于新疆古尔班通古特沙漠;它是流动沙丘的先锋物种,也能成功定居于人工固沙区(草方格)。对比研究了人工固沙区及自然流沙区的准噶尔无叶豆群落物种组成、种群密度、高度、盖度、生物量等特征及种群空间分布格局。结果表明:人工固沙区内的准噶尔无叶豆群落物种的科、属、种数均明显多于流沙区,而且2种生境物种相似性指数仅为0.522,表明人工固沙生境显著改变了群落物种组成与结构。除植株密度外,人工固沙区种群的高度,盖度,地上、地下及总生物量密度,单株地上、地下及总生物量等数量特征均显著高于流沙区。2种生境中准噶尔无叶豆种群空间分布的关联维数均接近2.0,体现了较强的个体空间相关性和空间占据能力。2种尺度(1 m和2 m)下的聚集度分析表明,2种生境中种群基本为随机分布。因而,人工固沙生境并未明显改变种群空间分布格局,这可能是种群的固有特性。人工固沙区的土壤有机质、全氮和有效氮含量显著高于流沙区,且与种群数量特征(植株密度除外)呈显著正相关。总之,与流沙区相比,人工固沙使地表得到固定,提高了土壤有机质和氮素含量,明显改变了定居于其中的准噶尔无叶豆种群主要数量特征,增强了种群的生存和适应能力。该结论对荒漠濒危植物的保育具有重要指示意义。     

苦瓜植株不同构件层次对支持物直径变化的行为反应

 采用实验生态学的方法,在提供不同直径支持物的情况下,对攀援植物苦瓜(Momordica charantia)不同构件层次的形态可塑性反应、生物量积累和生物量配置的变化进行了研究。结果表明：1)支持物直径对植株分枝出现的时机无明显影响,但分枝数量和分枝在主茎上的着生位置受支持物直径影响较大;2)植株个体水平比分枝水平对支持物直径的变化有更灵敏的反应,当支持物直径较大时,植株个体表现出对支持物较强的寻觅能力：主茎伸长受阻,比茎长变大,分枝数量和分枝率增大,分枝长度增加,比叶柄长度增加,根冠比减小;3)分枝形态特征和生物量特征对支持物直径的变化均无显著反应;4)苦瓜植株个体水平上的形态特征变化、生物量配置格局的改变以及分枝行为的变化是植株对支持物有效性变化的响应,有利于增强植株“寻觅”支持物的能力。     

风和风沙流对不同固沙植物生理状况的影响

在野外风洞条件下,对风和风沙流影响固沙植物的生理状况进行了研究。结果表明：风和风沙流对不同植物的影响不同,风对结皮和苔藓的影响在于吹蚀和干燥,对草本和灌木的影响主要是：１．通过迫使气孔关闭、降低叶温和胞间ＣＯ２浓度来抑制光合作用;２．加强蒸腾作用,减小水分利用率;３．降低叶片水势和土壤含水量,从而加强干燥作用。而风沙流对结皮和苔藓的影响主要在于剥蚀和堆积,对草本和灌木的影响主要是机械损伤作用,对光合和水分利用的影响将更加剧烈。不同类型植物对两种胁迫的适应程度不同。     

几种荒漠植物地上生物量估算的初步研究

在调查植物样方的基础上,利用植株冠幅特征如冠幅长与宽、株高、基径、新生枝条数、总枝条数等作为变量建立了估算典型荒漠植物地上生物量的模型,利用植株冠幅长与宽所建拟合方程对估算琵琶柴灌丛生物量的精确度较高,经验证,预测值与实测值的相关系数为O．9989,相对误差在4．79％～10．12％之间利用植株株高、基径所建拟合方程对估算梭梭、多枝柽柳生物量计算值与实测值的相关系数在0.9418以上,显著性检验表明相关系数均为极显著经验证,预测值与实测值的相关系数分别为0．9902和0．9875．相对误差分别为6．87％～19．22％和7．49％～18．47％。研究表明,在大面积荒漠植物生物量研究中,利用植株冠幅特征作为变量来估算琵琶柴灌丛地上生物量方法简便可行,用来估算梭梭、多枝柽柳等灌木地上生物量时存在一定误差     

固定沙丘结皮层藓类植物多样性及固沙作用研究

 根据对沙坡头固定沙丘生物结皮的物种调查、鉴定及盖度和生物量的研究,藓类植物共2科,7属,16种,以真藓（Bryum argenteum）为优势种。随着固沙时间的延长,生物结皮中藓类和藻类总盖度呈上升趋势;沿不同年代固定沙丘坡面向上等距离梯度,藓类盖度显著下降,而藻类盖度则平稳上升。对不同年代固定沙丘生物结皮优势种真藓的生物量和固沙量的测定表明,随沙丘固定年代的增加,真藓的生物量和固沙量增加,50年代固沙区结皮层真藓生物量是944.03 kg·hm-2,固沙量是3.925×104 kg·hm-2,但固沙率随时间的增加而减少。真藓的饱和吸水量与生物量呈正相关,50年代固定沙丘藓类结皮层可吸水7.06×103 kg·hm-2,说明藓类结皮层具有很强的吸水和保水能力,对于干燥少雨的沙漠地区具有十分重要的生态学意义。     

浑善达克沙地赖草分株种群与土壤资源异质性分析

对浑善达克沙地低湿滩地、滩地 -风沙沉积过渡区到风沙沉积区的赖草分株数、地上生物量、土壤水分和养分异质性进行了研究。结果表明 ,从低湿滩地、滩地 -风沙沉积过渡区到风沙沉积区 ,赖草的分株数和地上生物量与养分含量具有不同的变化趋势。赖草的分株数在 3种生境中都存在空间自相关 ,并且 ,从低湿滩地、过渡区到风沙沉积区空间自相关的变程逐渐增加。地上生物量同样存在空间自相关 ,但变程以过渡区最大 ,风沙沉积区最低。土壤水分仅仅在风沙沉积区存在空间自相关。在 3种生境条件下 ,土壤全氮和有机质的空间分布格局相似 ,都在低湿滩地和过渡区存在空间自相关且变程相似 ,在风沙沉积区不存在空间自相关。赖草的空间分布格局在低湿滩地为偏离随机适度聚集的分布格局 ,在过渡区近似于聚居分布 ,而在风沙沉积区为均匀分布格局     

细根与根系总质量比的相似性导致了共存的本土植物和入侵植物之间植物-土壤反馈的对比

三维土壤异质性对种子萌发影响的实验研究 土壤生物通过植物-土壤的反馈作用调控植物间相互作用和外来植物入侵。因此,探明植物-土壤反馈作用种间差异的形成原因,对于预测土壤生物在植物入侵过程中的作用具有重要价值。近期的研究发现,植物性状可以用于预测植物-土壤反馈作用。同时,研究发现植物入侵也与植物的一些性状相关联,暗示植物-土壤反馈作用通过植物性状与外来植物入侵之间存在关联,但尚缺乏实验证明。鉴于此,本研究选取了3对近缘入侵和本土植物为对象,比较了其植物-土壤反馈作用,探讨了植物-土壤反馈作用与植物根部性状的关联性。首先,通过种植实验植物3个月,驯化采自于田间的土壤。之后,将实验植物种植于对照和灭菌处理的同种或近缘(同科或同属)种驯化过的土壤中,评价同种或近缘种根际土壤生物对植物生长的净反馈作用(与灭菌土比较),并比较了两类土壤生物对植物的反馈作用。总体而言,同种或近缘种根际土壤生物对入侵与本土植物的净反馈作用无显著差异,两类土壤生物对入侵和本土植物的反馈作用亦无显著差异。土壤反馈作用的强度和种植于对照土壤中植物细根生物量比存在正相关关系,且入侵与本土植物细根生物量比无显著差异。这一发现表明：相似的细根生物量比可能是导致入侵与本土植物间土壤反馈作用无差异的一个重要原因。为提升人们对土壤生物在植物入侵过程中 作用的认识,亟需选取更多入侵与本土植物(尤其是亲缘关系较远的物种)开展实验研究,进一步探明植物性状、土壤反馈作用和外来植物入侵之间的关联性。     

科尔沁沙地南缘主要固沙植物旱季水分来源

探讨固沙植物水分来源以及物种间水分利用关系对揭示植物共存机理和固沙植被稳定机制具有重要意义.本研究选取科尔沁沙地南缘两种典型生境(固定沙丘和丘间低地)共12种固沙植物,通过测定植物水、同期降水、地下水和土壤水的稳定同位素比率(δD和δ¹⁸O),利用IsoSource模型计算植物对不同深度土壤水的利用比例,初步阐明半干旱沙区主要固沙植物旱季水分来源以及物种间的水分利用关系.结果表明:两种生境中不同生活型固沙植物水δD和δ¹⁸O差异显著,但丘间低地乔木和灌木差异不显著.在丘间低地从乔木到草本水分来源逐渐变浅,乔木和灌木主要利用50～150 cm或30～50 cm土壤水,半灌木主要利用10～30 cm土壤水,草本主要利用0～10 cm土壤水;固定沙丘灌木主要利用0～30 cm土壤水,半灌木则主要利用50 cm附近土壤水.表明旱季固定沙丘植物比丘间低地植物更依赖0～50 cm土壤水.固沙植物水分来源与植物生活型、根系分布范围有关,其中根系分布范围影响可能更大.     

Acquisition and allocation of resources in two waterlogging-tolerant grasses

This study focuses on the following questions: (i) whether reductions in root:shoot ratio have a cost in terms of nutrient balance of the plant, and (ii) whether changes in resource-allocation patterns are proportional among different resources. Our approach was to analyse the variations in the allocation pattern induced by soil waterlogging. A pot experiment was conducted to analyse the effects of waterlogging on biomass, phosphorus (P) and nitrogen (N) accumulation of Paspalum dilatatum and Danthonia montevidensis , two waterlogging-tolerant grasses. When changing from oxic to anoxic conditions, a common response of these and other waterlogging-tolerant grasses is a reduction in allocation to below-ground resources. It was observed that (i) the reduction in root:shoot ratio caused by waterlogging did not have a cost in terms of capacity for nutrient uptake; (ii) resource partitioning within aerial parts was less sensitive to treatments than partitioning between roots and shoots; and (iii) biomass does not appear to be a useful currency for evaluating nutrient-allocation patterns, as the allocation of P and N was inadequately represented by biomass. The results presented here indicate that the existence of compensation mechanisms reduces the predictive value of the partition of resources for the capacity of plants to acquire resources. Data on the allocation of nutrients in relation to biomass suggest that the assumptions of independence in the allocation pattern between biomass and limiting nutrients under the effects of environmental factors can be extended.     

Plant compensatory growth: a conquering strategy in plant–herbivore interactions?

We present a theoretical analysis that considers the phenotypic trait of compensatory growth ability in a context of population dynamics. Our model depicts a system of three interactors: herbivores and two different plant types referred to as ordinary and compensating. The compensating plant type has the ability to increase its intrinsic rate of biomass increase as a response to damage. This compensatory growth ability is maintained at the expense of a reduced growth rate in the absence of damage, where the ordinary plant type has the higher growth rate. Analysis of this system suggests that, even though a compensatory capacity of this kind will not imply an increase in equilibrium plant density, it will give a competitive advantage in relation to other plants, in the presence of a sufficiently efficient herbivore. Invasion of compensating plants into a population of non-compensating plants is facilitated by a high compensatory growth ability and a high intrinsic rate of plant biomass increase. Conversely, an ordinary plant can invade and outcompete a compensating plant when the herbivore is characterised by a relatively low attack rate, and/or when plant intrinsic growth rate is decreased.     

Morphological variation of Aechmea distichantha (Bromeliaceae) in a Chaco forest: habitat and size-related effects

Plants show different morphologies when growing in different habitats, but they also vary in their morphology with plant size. We examined differences in sun- and shade-grown plants of the bromeliad Aechmea distichantha with respect to relationships between plant size and variables related to plant architecture, biomass allocation and tank water dynamics. We selected vegetative plants from the understorey and from forest edges of a Chaco forest, encompassing the whole size range of this bromeliad. Plant biomass was positively correlated with most architectural variables and negatively correlated with most biomass allocation variables. Understorey plants were taller and had larger diameters, whereas sun plants had more leaves, larger sheath area, sheath biomass and sheath mass fraction. All tank water-related variables were positively correlated with plant biomass. Understorey plants had a greater projected leaf area, whereas sun plants had higher water content and evaporative area. Plasticity indices were higher for water-related than for allocation variables. In conclusion, there were architectural and biomass allocation differences between sun- and shade-grown plants along a size gradient, which, in turn, affected tank water-related variables.     

Plant and soil microbial responses to defoliation in temperate semi-natural grassland

There is much interest in understanding the nature of feedback mechanisms between plants and soil organisms in grazed ecosystems. In this study, we examine the effects of different intensities of defoliation on the growth of three dominant grass species, and observe how these plant responses relate to the biomass and activity of the microbial community in the root zone. Our data show that grassland plants with varying tolerances to grazing have markedly different growth responses to defoliation, and that these responses vary with the intensity of cutting. Defoliation of grasses which are tolerant to grazing, namely Festuca rubra and Cynosurus cristatus, leads to a reduction in root mass and an increase in the allocation of resources to shoots. In contrast, defoliation of a grass with low tolerance to grazing, Anthoxanthum odoratum, had little effect on root mass, but increased the relative allocation of resources below-ground. In all plant species, defoliation led to an increase in soil microbial biomass and C use efficiency in the root zone. This response was greatest in the root zone of A. odoratum and is likely to be related to changes in root exudation pattern following defoliation. The significance of these changes in relation to soil nutrient dynamics and plant nutrient uptake during regrowth require further exploration. This revised version was published online in June 2006 with corrections to the Cover Date.     

Functional aspects of root architecture and mycorrhizal inoculation with respect to nutrient uptake capacity

The aim of this research was to investigate the effect of arbuscular mycorrhizal (AM) colonisation on root morphology and nitrogen uptake capacity of carob ( Ceratonia siliqua L.) under high and low nutrient conditions. The experimental design was a factorial arrangement of presence/absence of mycorrhizal fungus inoculation ( Glomus intraradices) and high/low nutrient status. Percent AM colonisation, nitrate and ammonium uptake capacity, and nitrogen and phosphorus contents were determined in 3-month-old seedlings. Grayscale and colour images were used to study root morphology and topology, and to assess the relation between root pigmentation and physiological activities. AM colonisation lead to a higher allocation of biomass to white and yellow parts of the root. Inorganic nitrogen uptake capacity per unit root length and nitrogen content were greatest in AM colonised plants grown under low nutrient conditions. A better match was found between plant nitrogen content and biomass accumulation, than between plant phosphorus content and biomass accumulation. It is suggested that the increase in nutrient uptake capacity of AM colonised roots is dependent both on changes in root morphology and physiological uptake potential. This study contributes to an understanding of the role of AM fungi and root morphology in plant nutrient uptake and shows that AM colonisation improves the nitrogen nutrition of plants, mainly when growing at low levels of nutrients.     

沙蒿与油蒿灌丛的防风阻沙作用

沙蒿与油蒿广泛分布于我国沙漠地区,是沙地植被的重要建群种和优势种。在腾格里沙漠南缘半流动沙地,实测了两种典型固沙植物沙蒿与油蒿的防风阻沙作用,从灌丛空间构型对比分析了其防风阻沙机制。结果表明,沙蒿与油蒿灌丛均具有明显的降低风速作用,但油蒿灌丛较沙蒿灌丛具有更显著的防风作用,而且对灌丛后不同位置、近地面不同高度层风速的降低程度明显不同。在灌丛后6倍株高范围内,沙蒿灌丛对50cm高度风速降低程度显著大于20cm,而油蒿灌丛对近地面20cm高度层风速降低程度显著大于50cm。在相同风速下,油蒿灌丛后20cm高度平均风速是沙蒿灌丛的1/2,而50cm高度平均风速与沙蒿灌丛相近。同时,沙蒿灌丛阻沙作用弱,而油蒿灌丛具有明显的阻沙作用,单株积沙体积达到45.2±16.1dm3,积沙重量达到72.1±25.7kg,油蒿灌丛积沙量大小与灌丛结构间存在显著的正相关。研究表明,紧密型结构的油蒿灌丛是较松散型结构沙蒿灌丛更为理想的防风固沙植物,其灌丛分枝数多、分枝角度小、生物量大且多分布在近地面层是具有显著防风阻沙作用的根本原因,该结论可为干旱区防风固沙植被建设物种选择提供依据。     

Species traits and shoot-root biomass allocation in 20 dry-grassland species

Aims A plant has a limited amount of resources at any time and it allocates them to different structures. In spite of the large number of previous studies on allocation patterns within single species, knowledge of general patterns in species allocation is still very limited. This is because each study was done in different conditions using different methodology, making generalization difficult. We investigate intraspecific above- versus below-ground biomass allocation among individuals across a spectrum of dry-grassland plant species at two different developmental stages and ask whether allocation is age- and species specific, and whether differences among species can be explained by their life-history traits and phylogeny.Methods We collected data on above- and below-ground biomass of seedlings and adult plants of 20 species from a common garden experiment. We analysed data on shoot–root biomass allocation allometrically and studied the relationship between the allometric exponents (slopes on log–log scale), species life-history traits and phylogenetic distances.Important findings We found isometric as well as allometric patterns of biomass allocation in the studied species. Seedlings and adult individuals of more than half of the species differed in their above- versus below-ground biomass allometric exponents. Seedlings and adult individuals of the remaining species differed in their allometric coefficients (intercepts). Annual species generally allocated proportionally more to above- than below-ground biomass as seedlings than as adults, whereas perennial species showed the opposite pattern. Plant life-history traits, such as plant life span, age of first flowering, month in which the species begin flowering and specific leaf area were much more important in explaining differences in shoot–root allometry among species than were phylogenetic relationships. This suggests that allocation patterns vary greatly among closely related species but can be predicted based on species life-history traits.     

腾格里沙漠人工植被区固沙灌木影响深层土壤水分的动态模拟研究

灌木是我国沙漠地区主要的优势植物类型,固沙灌木的存在有益于沙丘的固定进而有利于退化沙漠生态系统的生态恢复。基于腾格里沙漠沙坡头地区50多年的人工植被区长期观测研究,建立生态-水文模型模拟了该地区固沙灌木盖度和深层土壤水分的动态变化过程,结果表明,固沙灌木的建立改变了沙区原有的生态-水文过程,在固沙灌木建立40多年后,固沙灌木的盖度和深层土壤水分达到了新的平衡状态。灌木盖度逐渐稳定在(10±0.9)%左右,而深层土壤水分稳定在(2.58±0.2)%左右。因此,在年均降雨量为186mm的腾格里沙漠沙坡头地区,土壤水分的最大植被承载力为:灌木和生物土壤结皮的盖度分别维持在10%和60%,深层土壤水分维持在3%左右。     

Differences in allocation patterns in clonal and sexual offspring in a woodland pseudo-annual

We compared the patterns of allocation to reproduction among seed-derived and clonal offspring of a woodland pseudo-annual. Pseudo-annuals are clonal plants which survive the winter only as seeds and hibernacles produced by the rhizome system. Previous studies indicate that flowering is related to the size of these hibernacles. Since seedlings do not have a hibernacle, we did not expect that these plants would reproduce sexually. Assuming a trade-off between sexual and asexual reproduction, and assuming a linear relationship between vegetative plant weight and weight of all reproductive structures (i.e., rhizomes, hibernacles, inflorescences, and seeds), we expected that seed-derived plants would have a stronger biomass allocation to rhizomes and hibernacles. Since resource supply affects plant size, and thus hibernacle and seed production, we also subjected the plants to different levels of shade. At the start of the experiment seed-derived and clonal offspring hardly differed in total fresh weight. At the final harvest in September seed-derived and clonal offspring did not differ in vegetative plant weight (i.e., leaves, stems, and roots). Only light availability significantly affected these plant structures. As predicted, seed-derived plants did not flower in either of the light treatments. Seed-derived plants allocated more biomass to rhizomes and hibernacles, but this was only significant in the highest-light treatment. This result was due only to an increase in the number of hibernacles. Dry weight of single hibernacles was not affected by plant type. The ecological implications of this allocation pattern are discussed. Received: 2 October 1997 / Accepted: 8 March 1998     

黄土高原蓝藻和藓类生物结皮对草本植物多样性及生物量的影响

生物结皮在干旱和半干旱气候区占据了广阔的生态位，深刻影响草本植物的空间格局，但其影响程度和作用途径目前仍存有争议。分别以黄土高原风沙土和黄绵土上的蓝藻结皮和藓结皮为对象，以无结皮为对照，通过样方法调查并测定了生物结皮与无结皮样地中草本植物的多样性和生物量，继而结合灰色关联分析和结构方程模型，对比分析了两类生物结皮对草本植物多样性和生物量的影响程度及作用途径。结果表明：（1）风沙土上，无结皮样地的优势科为禾本科和菊科，蓝藻结皮样地的优势科为藜科，藓结皮样地的优势科为禾本科，而黄绵土上无结皮样地的优势科为菊科、禾本科和豆科，蓝藻结皮样地的优势科与无结皮相似，藓结皮样地的优势科为禾本科和菊科；（2）与无结皮相比，风沙土上蓝藻结皮和藓结皮样地中多年生草本的株数分别增加了30%和16%，而黄绵土上两类生物结皮样地中多年生草本株数则分别增加了44%和减少了46%；（3）与无结皮相比，风沙土上蓝藻结皮和藓结皮样地中草本植物的Patrick和Shannon指数分别降低了1.0、0.2和3.4、0.5；而黄绵土上蓝藻结皮和藓结皮样地中两种多样性指数分别降低了4.8、0.2和4.2、0.4；（4）风沙土上蓝藻结皮和藓结皮样地的草本植物生物量比无结皮减少了111.5 g/m²和145.0 g/m²，而在黄绵土上则分别减少了127.8 g/m²和172.2 g/m²；（5）灰色关联分析显示，土壤穿透阻力对草本植物的多样性影响最大（关联系数分别为0.960和0.996），而NH₄⁺-N和NO₃^--N含量对草本植物的生物量影响最大（两种土壤上关联系数分别为0.727和0.752）。综上，黄土高原生物结皮显著降低了草本植物的多样性和生物量，其途径为通过增加土壤穿透阻力以降低草本植物的种子萌发几率并限制幼苗根系伸展，同时通过土壤氮竞争限制了草本植物生长。