植物地上竞争与地下竞争研究进展

植物地上光竞争和地下水分与营养的竞争过程十分复杂,并且与竞争机制密切相关。从地上和地下竞争理论、两种竞争过程的相互作用机制以及分离地上和地下竞争的研究方法和表达指标等几个方面综述和分析目前的研究状况,以期为国内研究者在本领域进一步展开实验提供理论依据和实验设计参考。总结了与地上、地下相对竞争强度有关的优化分配理论,地上、地下竞争随生产力梯度的变化,资源异质性对地上、地下竞争机制的影响。介绍了地上竞争和地下竞争相互作用类型以及目前提出的两种作用机制。对常用的3种分离地上、地下竞争的方法:盆分隔、间植分隔和目标植分隔法以及表达竞争强度和竞争重要性的指标和公式进行了归纳。提出未来的研究内容,认为在开展此类研究时,应考虑到实验植物的生理学特性、发育状况、立地生产力跨度范围以及竞争持续时间等方面因素,并认为发展地下竞争研究、深入探讨根系间相互作用的影响因子和过程是当前的研究热点。     

种内竞争与个体大小不等性

种内竞争的后果是生态学家一直关注的问题。除了影响种群产量以及个体的平均大小,种内竞争还会改变种群内个体大小的差异。理解种内竞争对个体大小不等性的影响对于认识种内竞争的进化生物学后果非常重要。介绍了相关的理论和实验研究案例．这些工作验证了种内竞争扩大了植株大小不等性,也区分了对称性竞争和非对称性竞争对其的影响。这些案例对于丰富中学生物学教学和兴趣类活动课有很好的参考价值。     

利用竞争指数评价水曲柳落叶松种内种间空间竞争关系

林木之间的竞争包括地上和地下两部分。竞争影响空间的确定对预测生长有重要的意义。本研究将竞争影响空间划分为两部分：距对象本最近的一圈,称为内圈,包括地上和地下竞争影响;除最近一圈以外称为外圈,主要为地下竞争影响。在此基础上将竞争指数分解为不同的竞争指数分量。并以水曲柳和落叶松纯林和混交林为例,定量地分析了竞争影响空间内地上竞争和地下竞争,种内和种间竞争影响大小。结果表明,纯林中两树体最近一圈内地上和     

地上竞争与地下竞争对科尔沁沙地榆树幼苗生长的影响

榆树疏林草原对科尔沁沙地植被恢复和景观保护有着重要意义.本文采用双因素两水平控制试验,从幼苗生物量、地下/地上生物量、茎高、根茎比、叶片数等方面,研究了草-树地上、地下竞争对科尔沁沙地榆树幼苗生长的影响.结果表明:对于1年生榆树幼苗,单株平均生物量表现为无竞争＞地上竞争＞全竞争＞地下竞争;地下/地上生物量表现为地下竞争＞全竞争＞无竞争＞地上竞争;幼苗高度表现为地上竞争＞无竞争＞全竞争＞地下竞争;根茎比表现为地下竞争＞全竞争＞无竞争＞地上竞争;叶片数表现为地上竞争＞无竞争＞地下竞争＞全竞争.地下竞争对1年生榆树幼苗生长影响显著,而地上竞争对榆树幼苗生长无显著影响.地上竞争与地下竞争对2年生榆树幼苗生长的影响均不显著.科尔沁沙地草本植物对榆树幼苗生长的影响主要通过地下竞争的方式实现,但地下竞争并没有改变榆树幼苗的资源分配方式.随榆树幼苗龄级的增长,草本植物竞争作用的影响逐渐减弱.     

植物竞争对3种移植树苗生长的影响

在植物幼苗生长过程中,总是受到包括地下根竞争在内的各种竞争影响。植物间的竞争主要同有效光辐射、水分和各种营养相关。当外来植物侵入森林群落时,可能受到群落中其它植物竞争的影响。该文通过移植尾叶桉(Eucalyptus urophylla)、大叶相思(Acacia auriculaeformis)两种外来种和本地种荷木(Schima superba)幼苗,挖沟排除根竞争和砍树创造林窗来排除地上竞争的野外实验,研究植物竞争对幼苗生长的影响。有根竞争时,荷木、尾叶桉和大叶相思幼苗的生物量和净初级生产力均小于没有根竞争时,可见地下根竞争对3种幼苗生长有抑制性影响。尾叶桉、大叶相思和荷木3种幼苗受到的总竞争强度分别是0.357 9、0.338 3和0.198 9,受到的地下根竞争强度则分别是0.104 3、0.053 04和0.118 8,受到的地上竞争强度则分别是0.285 1、0.277 0和0.090 85。尾叶桉和大叶相思两种幼苗受到的总竞争强度间的差异不显著,但都显著大于荷木;3种幼苗受到的地上竞争强度间的差异同总竞争强度情况相似;尾叶桉和荷木两种幼苗受到的地下根竞争强度间的差异不显著,但都显著大于大叶相思。地上竞争对阳性树种尾叶桉和大叶相思两种幼苗的生长影响大,而地下根竞争则对耐阴性强的荷木幼苗影响大。尾叶桉和大叶相思两种外来种幼苗受到的总竞争强度均大于本地种荷木幼苗,这反映了这两种外来树种侵入次生林这样的群落受到竞争影响大。     

水曲柳苗木地下竞争与地上竞争的定量研究

对3种密度进行栽培试验,利用通径分析的方法,研究了水曲柳地下竞争和地上竞争的关系及对总竞争的影响。结果表明,水曲柳苗木的地下部分生物量、地上部分生物量和总生物量与营养空间有密切关系。随着苗木空间距离增加,由生物量计算的竞争指数下降。在同一密度条件下,地下竞争指数明显大于地上竞争指数。由于地下生长与地上生长的相互作用,各竞争指数之间具有明显的相关性。但是地下竞争和地上竞争对总竞争的影响是不同的,通径分析可以定量的区分地下竞争和地上竞争的相对大小。地下竞争对总竞争的直接作用范围在0．5543～0．7426之间,明显大于地上竞争对总竞争的直接作用(0．2851～0．5282)。随着距离的增加,单株苗木的生长空间加大,地上部分的竞争作用增加,地下部分的竞争程度减弱。但是,地下根系的竞争在水曲柳苗木总的竞争中占有重要地位。     

湿热河谷芭蕉群落优势物种空间分布格局与竞争对称性研究

竞争关系的对称性是塑造植物物种空间分布格局和影响物种共存重要机制之一,对其开展研究不仅对于揭示植物群落构建机制具有重要意义,也有助于解析群落演替的驱动力。在贵州赤水湿热河谷内的芭蕉群落中建设固定样地,对1989棵植物个体进行空间定位和物种识别并记录胸径等个体属性。采用对数相关函数和标记变异系数分析（marked variogram）等空间统计方法检测优势种的空间分布格局和竞争对称性,结果如下：（1）芭蕉（Musa sp.）,粗糠柴（M.philippensis）,粗叶木（L.chinensis）,川钓樟（L.pulcherrima）,红果黄肉楠（A.cupularis）等5种优势种均表现为小尺度下的种内聚集分布,其中芭蕉主要采用克隆繁殖,其重要值达33.6%,且种内聚集度最大。（2）芭蕉种群在5 m尺度下呈现出显著的种内对称性竞争,这一竞争过程减少了种群自疏作用并推动种内聚集分布格局的形成,增加了该种群在森林中的生存力。（3）芭蕉与其余4种优势种之间的不对称竞争过程导致了种间负关联。结果表明,不同优势种在种间竞争对称性上的差异说明了植物种群繁殖和扩散方式是群落构建的重要影响因素之一,相比于有性繁殖过程,采用无性克隆繁殖的植物个体可通过种内对称竞争机制来减少自疏以增加种群生存力,并通过与相邻异种个体的不对称竞争过程来提升种群获取有限资源的能力,进而成为演替中的优势个体。研究结果可为今后制定植物物种多样性保护政策提供科学参考。     

湿地植物间竞争和促进互作的研究进展

湿地植物间关系主要包括竞争(负效应)和促进(正效应),是湿地生态研究的核心内容之一,对植物分布、群落和生态系统组成具有重要的调节作用。本文阐述了竞争理论及影响湿地植物竞争能力的主要因素,介绍了促进作用及其在湿地生态系统中的形成机理,详细综述了植物间关系转变的主要理论以及影响湿地植物间关系转变的主要环境因子(水位、温度、盐度、生物因子)。随着环境胁迫强度的增加,植物间关系一般会从竞争转变为促进作用,即优越条件下以竞争为主,胁迫条件下以促进作用为主。针对当前湿地植物间关系的研究多集中于滨海盐生湿地,类型较单一的情况,认为将来需加强对淡水湿地、河口湿地等各类系统的研究。同时,加强湿地植物间关系转变的生物学机制、时间效应,对新研究方法的探索等方面的研究也有助于进一步理解湿地植物间竞争和促进作用的发生及转变机理。     

林窗中植物竞争强度随林窗发育的变化

 通过野外移植外来种尾叶桉(Eucalyptus urophylla)、大叶相思(Acacia auriculaef ormis)和本地种荷木(Schima superba) 3种幼苗,挖沟排除根竞争和砍树创造林窗排除地上竞争的实验,来测定林窗形成后植物地下竞争和地上竞争强度的变化以及这些竞争对3种幼苗生长的影响。在林窗内,尾叶桉、大叶相思和荷木受到的地下竞争强度和地上竞争强度均呈现出随林窗年龄的增长而逐渐增大的趋势。外来种尾叶桉和大叶相思受到的地上竞争强度大于本地种荷木受到的地上竞争强度,而荷木受到的地下竞争强度大于两种外来种受到的地下竞争强度。在只有地下竞争和地上竞争时,这3种幼苗的相对生长速率均小于没有竞争时,这反映了地下竞争和地上竞争对3种幼苗的生长有抑制性影响,而且这种影响在本地种和外来种间存在差异。     

不同栽植方式下紫椴幼苗生物量分配及资源利用分析

 通过紫椴（Tilia amurensis Rupr.）幼苗在不同植距下纯栽和与落叶松(Larix gmelini)的混栽沙培试验,初次应用通径分析,研究紫椴幼苗生物量分配和资源利用机制,分析种内和种间竞争及其模式。研究发现总体上植距增大有利于紫椴根、茎、叶各器官的生长。通过通径分析说明,紫椴地下根系对总生长的贡献为0.236 1～0.286 9,地上茎、叶对总生长的贡献为0.732 5～0.775 8,地上部分的贡献是地下根系的2.95倍。表明紫椴幼苗对光具有较强的需求和竞争能力。紫椴和落叶松之间存在悬殊较大的生物量差距,落叶松的根、茎、叶生物量约是相应混栽紫椴的2、5、4倍。通过比较纯栽和混栽试验中的直接通径系数,与落叶松混栽后,紫椴对地下水分养分的竞争下降36.84%,对地上光照空间的竞争增大11.63%。随植距减小落叶松对紫椴的影响增大,紫椴根系生长受落叶松种间竞争影响不显著;地上茎、叶生长只在一定植距范围内受落叶松影响。紫椴与落叶松种间竞争是不同资源利用等级的两树种间的相互作用,以对称性地下竞争与落叶松“平等”地占有地下水分养分资源,从而受落叶松地下竞争较小;在一定空间范围内落叶松以非对称性竞争优先占有地上光照资源影响紫椴生长。     

Biodiversity and belowground interactions mediate community invasion resistance against a tall herb invader

Aims Species-rich plant communities are hypothesized to be more resistant against plant invasions because they use resources in a more efficient way. However, the relative contributions of aboveground competition and belowground interactions for invasion resistance are still poorly understood.Methods We compared the performance of Knautia arvensis transplants growing in plots differing in plant diversity both under full competition and with shoots of neighbors tied back to determine the relative strength of aboveground competition in suppressing this test invader without the confounding effect of shading. In addition, we assessed the effects of belowground competition and soil-borne pathogens on transplant performance.Important findings Both aboveground competition and plant species richness strongly and independently affected invader performance. Aboveground biomass, height, leaf mass per area and flowering of transplanted individuals of K. arvensis decreased with increasing species richness of the host community. Species-rich and species-poor communities both imposed equally strong aboveground competition on K. arvensis. However, belowground interactions (especially belowground root competition) had strong negative effects on transplant performance. In addition, the presence of grasses in a plant community further reduced the performance of K. arvensis. Our results suggest that belowground competition can render species-rich host communities more suppressive to newly arriving species, thus enhancing community invasion resistance.     

禾-豆混播草地中土壤水分与种间关系研究进展

从植物形态变化和生理反应两方面探讨了禾草和豆科牧草在干旱环境下所采取的不同生存策略,阐述了禾-豆之间存在的水分竞争关系以及豆科植物的水力提升作用对水分资源高效利用的意义,分析了干旱对豆科植物固氮和转氮能力的影响.提出应探讨适于我国干旱区域的深根系豆科牧草与禾草的共存机制,认为揭示水分因子对地上竞争和地下竞争以及对两种竞争间反馈关系的影响是研究其共存机制的一个重要方面.     

Plant life history and above–belowground interactions: missing links

The importance of above–belowground interactions for plant growth and community dynamics became clear in the last decades, whereas the numerous studies on plant life history improved our knowledge on eco‐evolutionary dynamics. However, surprisingly few studies have linked both research fields despite their potential to increase our mechanistic understanding of how above belowground interactions are governed. Here I briefly review studies on above–belowground interactions and plant life history and identify important research gaps. To advance our understanding of ecological strategies and eco‐evolutionary dynamics of plants and their associated organisms it is warranted to elucidate the interconnectivity and tradeoffs of plant life history traits of growth, defence, reproduction, nutrient cycling and the functional composition of above‐ and belowground heterotrophic communities. Using the concept of tradeoffs in growth, reproduction and defence we can postulate that plants in rich soil grow, reproduce and die fast whilst avoiding above‐ and belowground antagonists, whereas plants in poor soil grow slow, live and reproduce longer and invest in above‐ and belowground mutualists and defences. However, alternative scenarios are possible and depend on the selection pressure by above‐ and belowground mutualists and antagonists during plant ontogeny and via after‐life effects. To elucidate missing links between life history traits and above–belowground interactions, complementary modelling and empirical studies are needed that reveal the coupling between below‐ and aboveground plant traits of growth, defence and reproduction, their heritability and their cost/benefit relation. These cost/benefit analyses of defence should span from individuals to future generations, taking feedback effects via altered biotic communities and resource competition into account. The role of soil fertility in steering plant life history traits requires explicit testing of trans‐generational trait shifts in growth, defence, reproduction, cost/benefit of associations with mutualists and antagonists and soil feedbacks across plant genotypes/species with distinct life history traits, grown across soil fertility gradients.     

Interactions between spatially separated herbivores indirectly alter plant diversity

Above‐ and belowground herbivores promote plant diversity when selectively feeding on dominant plant species, but little is known about their combined effects. Using a model system, we show that neutral effects of an aboveground herbivore and positive effects of a belowground herbivore on plant diversity became profoundly negative when adding these herbivores in combination. The non‐additive effects were explained by differences in plant preference between the aboveground‐ and the belowground herbivores and their consequences for indirect interactions among plant species. Simultaneous exposure to aboveground‐ and belowground herbivores led to plant communities being dominated by a few highly abundant species. As above‐ and belowground invertebrate herbivores generally differ in their mobility and local distribution patterns, our results strongly suggest that aboveground–belowground interactions contribute to local spatial heterogeneity of diversity patterns within plant communities.     

Hierarchical plant responses and diversity loss after nitrogen addition: testing three functionally-based hypotheses in the Inner Mongolia grassland

Background

Numerous studies have shown that nitrogen (N) deposition decreases biodiversity in terrestrial ecosystems. To explain the N-induced species loss, three functionally based hypotheses have been proposed: the aboveground competition hypothesis, the belowground competition hypothesis, and the total competition hypothesis. However, none of them is supported sufficiently by field experiments. A main challenge to testing these hypotheses is to ascertain the role of shoot and root competition in controlling plant responses to N enrichment. Simultaneously examining both aboveground and belowground responses in natural ecosystems is logistically complex, and has rarely been done.

Methodology/Principal Findings

In a two-year N addition experiment conducted in a natural grassland ecosystem, we investigated both above- and belowground responses of plants at the individual, species, and community levels. Plants differed significantly in their responses to N addition across the different organizational levels. The community-level species loss was mainly due to the loss of perennial grasses and forbs, while the relative abundance of plant species was dependent mainly on individual-level responses. Plasticity in biomass allocation was much smaller within a species than between species, providing a biological basis for explaining the functionally based species loss. All species increased biomass allocation to aboveground parts, but species with high belowground allocations were replaced by those with high aboveground allocations, indicating that the increased aboveground competition was the key process responsible for the observed diversity loss after N addition in this grassland ecosystem.

Conclusions/Significance

Our findings shed new light on the validity of the three competing hypotheses concerning species loss in response to N enrichment. They also have important implications for predicting the future impacts of N deposition on the structure and functioning of terrestrial ecosystems. In addition, we have developed a new technique for ascertaining the roles of aboveground and belowground competition in determining plant responses to N fertilization.     

Empirical and theoretical challenges in aboveground–belowground ecology

A growing body of evidence shows that aboveground and belowground communities and processes are intrinsically linked, and that feedbacks between these subsystems have important implications for community structure and ecosystem functioning. Almost all studies on this topic have been carried out from an empirical perspective and in specific ecological settings or contexts. Belowground interactions operate at different spatial and temporal scales. Due to the relatively low mobility and high survival of organisms in the soil, plants have longer lasting legacy effects belowground than aboveground. Our current challenge is to understand how aboveground–belowground biotic interactions operate across spatial and temporal scales, and how they depend on, as well as influence, the abiotic environment. Because empirical capacities are too limited to explore all possible combinations of interactions and environmental settings, we explore where and how they can be supported by theoretical approaches to develop testable predictions and to generalise empirical results. We review four key areas where a combined aboveground–belowground approach offers perspectives for enhancing ecological understanding, namely succession, agro-ecosystems, biological invasions and global change impacts on ecosystems. In plant succession, differences in scales between aboveground and belowground biota, as well as between species interactions and ecosystem processes, have important implications for the rate and direction of community change. Aboveground as well as belowground interactions either enhance or reduce rates of plant species replacement. Moreover, the outcomes of the interactions depend on abiotic conditions and plant life history characteristics, which may vary with successional position. We exemplify where translation of the current conceptual succession models into more predictive models can help targeting empirical studies and generalising their results. Then, we discuss how understanding succession may help to enhance managing arable crops, grasslands and invasive plants, as well as provide insights into the effects of global change on community re-organisation and ecosystem processes.     

高寒草地植物群落地上-地下净初级生产力权衡

植物生产力分配和权衡是植物生态学研究中的热点,反映植物对环境的适应性,是了解植物响应全球气候变化的关键。青藏高原作为气候变化敏感区,研究其植物地上与地下部分权衡对了解高寒草地植被生存策略和生态系统可持续发展具有重要意义。目前,生物量分配调控机制已被广泛研究,但主要使用植物根冠比和地上-地下生物量比等方法来表征植物分配模式,缺乏考虑因植物生长周期导致的差异。使用青藏高原高寒草地103个样点的地上和地下净初级生产力数据,分析高寒草甸和高寒草原两种主要草地类型的地上-地下净初级生产力权衡关系。利用气候因素和土壤因素等相关数据,结合方差分析、相关分析、相对重要性分析和结构方程模型的方法,探究环境因素对两种草地类型地上-地下净初级生产力权衡的影响机制。研究发现：（1）高寒草甸的地上净初级生产力、地下净初级生产力和土壤养分含量显著高于高寒草原（P<0.05）;（2）高寒草地植被生产力均向地下权衡（0.0199）,且高寒草原（0.0354）的权衡值高于高寒草甸（0.0173）;（3）结构方程模型发现,年平均降水量、土壤容重和土壤速效氮含量是影响高寒草甸生产力权衡的主导因子,而年平均温度和年平均降水量是影响高寒草原生产力权衡的主导因子。研究表明高寒草甸的生产力权衡主要受气候和土壤因素共同影响,而高寒草原主要受气候因素调节。研究为理解植物地上-地下生物量分配调控机制提供了新的视角和方法,对系统了解高寒草地生物量分配模式和准确预测高寒草地植被动态过程具有指导意义。     

Comparative impacts of aboveground and belowground enemies on an invasive thistle

Most research examining how herbivores and pathogens affect performance of invasive plants focuses on aboveground interactions. Although important, the role of belowground communities remains poorly understood, and the relative impact of aboveground and belowground interactions is still debated. As well, most studies of belowground interactions have been carried out in controlled environments, so little is known about the role of these interactions under natural conditions or how these relationships may change across a plant's range. Using the invasive plant Cirsium arvense, we performed a reciprocal transplant experiment to test the relative impacts of above‐ and belowground interactions at three sites across a 509‐km latitudinal gradient in its invaded range in Ontario, Canada. At each site, C. arvense seedlings were protected with above‐ and/or belowground exclosures in a factorial design. Plant performance (biomass, height, stem thickness, number of leaves, length of longest leaf, maximum rhizome length) was greatest when both above‐ and belowground exclosures were applied and lowest when no exclosures were applied. When only one type of exclosure was applied, biomass generally improved more with belowground exclosures than with aboveground exclosures. Despite site‐to‐site differences in foliar damage, root damage, and mesofaunal populations, belowground interactions generally had a greater negative impact on performance than aboveground herbivory alone. These results stress the importance of including both aboveground enemy interactions and plant–soil interactions in studies of plant community dynamics and invader performance.     

Linking aboveground and belowground diversity

Aboveground and belowground species interactions drive ecosystem properties at the local scale, but it is unclear how these relationships scale-up to regional and global scales. Here, we discuss our current knowledge of aboveground and belowground diversity links from a global to a local scale. Global diversity peaks towards the Equator for large, aboveground organisms, but not for small (mainly belowground) organisms, suggesting that there are size-related biodiversity gradients in global aboveground-belowground linkages. The generalization of aboveground-belowground diversity relationships, and their role in ecosystem functioning, requires surveys at scales that are relevant to the organisms and ecosystem properties. Habitat sizes and diversity gradients can differ significantly between aboveground and belowground organisms and between ecosystems. These gradients in biodiversity and plant community trait perception need to be acknowledged when studying aboveground-belowground biodiversity linkages.