植物对称性竞争与非对称性竞争研究进展及展望

植物个体间的竞争是植物种群和群落中的普遍现象, 一直是生态学家关注的焦点, 尤其是植物在生长过程中与邻体的地上竞争。在环境水分胁迫或营养胁迫条件下, 植物根系间的地下竞争同样重要。然而, 有关植物体地上、地下竞争模式的相关理论一直存在分歧和争论。该文扼要介绍了植物地上、地下竞争的概念及模式, 简要阐述了植物对称性竞争与非对称性竞争相关研究的理论模型及其检验方法, 概述了影响植物间竞争甚至改变竞争模式的主要因素及其意义, 讨论了研究物种对称性竞争和非对称性竞争的模式和机制对生态学其他研究领域的影响和重要意义, 并对植物对称性竞争与非对称性竞争的相关研究进行了展望。     

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。     

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。     

苗期无瓣海桑与互花米草的营养竞争关系研究

采用间植分隔法开展了无瓣海桑和互花米草营养竞争关系研究,结果表明竞争作用刺激了无瓣海桑高、径、冠幅、分枝及叶片数的增长,却抑制了叶面积的增加。地上竞争可增加无瓣海桑的地上生物量,但对根冠比影响不大。地下竞争增加了互花米草的地上生物量,降低了其根生物量,此时根冠比最小。地下竞争促进了无瓣海桑的主根生长,抑制了侧根生长,而地上竞争则正相反;随着竞争程度的加深,根活力逐渐降低。地下竞争时互花米草根活力显著下降。无瓣海桑和互花米草在有地上竞争时叶绿素a和b的含量均降低。各竞争处理未对P、K、Ca和Mg元素的含量产生显著影响,但地上竞争显著降低了N元素在无瓣海桑和互花米草根茎叶中的含量。互花米草的竞争强度略高于无瓣海桑,但是地上竞争强度无瓣海桑远远高于互花米草,而二者地下竞争强度相当。     

根系间的相互作用——竞争与互利

植物根系间的相互作用分为竞争和互利两种形式 ,它是决定植物群落动态变化和群落结构的重要因素。根系间的竞争包括植株个体自身根系的竞争以及个体与个体根系间 (同种或异种 )的竞争两方面 ,前者的发生在农林系统中是不可避免的并且很难调控 ,后者可以分为种间植物根系的竞争和种内植物根系的竞争。还阐述了根系的竞争能力和与其密切相关的根系生长率、根组织的新陈代谢、植物的生长形式和根系的空间结构等植物特性 ,同时介绍了根系对水和养分的竞争机理、形式、影响以及竞争强度计算方法。接着具体分析包括根系错位在内的各种根系互利现象和相关机理。影响根系间相互作用的限制性因子有土壤营养的异质性、大气 CO2 浓度、地下草食生物、根系生产力和生物量、根系结构、形态和生理调节、土壤养分的扩散性以及植物间距等。随着科技的进步和各门学科的发展 ,未来根系的研究方向主要体现在结合实践优化农林系统中不同物种间的作用关系、预测根系竞争在全球气候变化下的发展规律、更新实验研究方法及手段研究作用机理等 3个方面。     

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。     

高寒草甸植物群落物种多样性和生产力关系的光竞争研究

通过施肥形成的生产力由低到高的过程中,物种多样性往往降低。总体竞争假说认为对所有资源的竞争作用对多样性的影响随着生产力提高而加剧,导致物种多样性的下降;光竞争假说则认为随着生产力提高,种间竞争从低生产力时的地下竞争转向高生产力时的光竞争,是光竞争导致了物种多样性的下降。为了验证这两种假说,本文通过在甘南玛曲高寒草甸的均匀施肥实验,研究了光竞争对高寒草甸植物群落物种多样性和生产力关系的影响。结果表明：(1)随着施肥梯度的增加,大部分植物的生长速率加快,高度和叶面积增加;(2)随着施肥梯度的增加,植物群落地上总的生物量提高,叶面积指数增加,透光率降低,物种多样性减少;(3)个体大小不对称的光竞争导致了高寒草甸植物群落物种多样性随施肥梯度的增加而减少。     

关于植物竞争强度CI测定方法的评述

关于竞争强度(ｃｏｍｐｅｔｉｔｉｏｎｉｎｔｅｎｓｉｔｙ,ＣＩ)与环境条件相关的假设,存在于一些植物群落的组织结构模型中。根据“ＲＣＳ(杂草竞争耐压)模型”[10]的假设,植物的竞争强度随生境生产力增大而增大;相反,“ＲＲ(资源利用率,ＲｅｓｏｕｒｃｅＲａｔｉｏ)模型”[17,19]则假设植物地上部分与地下部分的比率虽然存在竞争差异,但总的竞争强度不受生境生产力的影响;最后,植物群落的“ＨＴ(生境模块,ＨａｂｉｔａｔＴｅｍｐｌｅｔ)模型”[14],假设竞争强度与生境生产力不相关,但受干扰强度控制。最近的研究指出,用于量化竞争强度的精…     

种间竞争对香蒲与芦苇生长的影响

为了探究种间竞争对香蒲（Typha domingensis）与芦苇（Phragmites australis）生长的影响,利用根系分隔盆栽试验研究了3种不同分隔方式条件下香蒲与芦苇的种间竞争特性,包括塑料膜分隔（根系完全分隔,无相互作用,无物质交换）、尼龙网分隔（根系部分分隔,无相互作用,有物质交换）和不分隔（根系完全相互作用,有物质交换）,分析了香蒲与芦苇根系形态和地上部生长的变化,探究香蒲与芦苇种间竞争的差异。结果发现（1）在尼龙网分隔和不分隔处理中芦苇具有明显的劣势。与塑料膜分隔处理相比,芦苇的总生物量、植株密度在尼龙网分隔和不分隔处理中分别减少了39.14%、49.41%和82.08%、79.22%,总根长、总根表面积、总根体积分别减少了40.53%、44.84%、62.52%和85.7%、82.45%、89.67%,且均具有极显著差异（P<0.01）;根系分隔方式也影响芦苇的株高、茎粗和叶片数,表现为不分隔 > 塑料膜分隔 > 尼龙网分隔。（2）与塑料膜分隔处理相比,香蒲总生物量在尼龙网分隔和不分隔中虽有增加,但差异不显著,植株密度和株高在尼龙网分隔和不分隔处理中都有增加且具有显著差异（P<0.05）,其总根长、总根表面积、总根体积在尼龙网分隔中分别增加了57.93%、26.5%、8.04%,但在不分隔处理中分别减少了11.57%、14.92%、11.39%（P<0.05）,虽然根系的相互作用对香蒲根系的生长具有促进作用,但植物种间根系相互作用越强,对两者的生长越不利。（3）在不同的分隔方式中,芦苇与香蒲间也存在明显变化。在不分隔处理中,香蒲的生物量和植株密度是芦苇的1.7倍和6.74倍,与塑料膜分隔处理相比增加了6倍,表明芦苇与香蒲根系的完全相互作用,显著削减了芦苇的繁殖生长,增加了香蒲的根系分蘖。（4）通过种间相互作用强度（RII值）分析也表明,尼龙网分隔和不分隔处理下芦苇表现为抑制作用（RII值为负值）,香蒲表现为促进作用（RII值为正值）。香蒲与芦苇互作对芦苇具有抑制作用,说明种间相互作用是能改变植物的适应性和植物群落的繁殖,同时也表明植物根系不仅在吸收土壤中的水和养分中起着关键作用,在种间关系中也起着重要作用。因此利用种间竞争控制植物生长,可以为保护生物多样性和生态系统的功能提供有效的技术支撑。     

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     

The separation of above- and below-ground competition in plants A review and critique of methodology

The relative merits of different methods for separating root and shoot competition in plants are reviewed. The `Divided Pot' technique involves vertical partitions that divide above- and below-ground competition within pots. This design usually creates unrealistic competition for light by using artificial barriers around pot perimeters to contain above-ground growth and by using a constant orientation of incident light from directly above. The `Row' technique involves parallel rows of plants that are separated by vertical partitions above- and below-ground. Although, this design may be well suited to some agricultural applications, its value for field studies of wild plants is limited. The Divided Pot and Row techniques have been associated often with the replacement series design, which has a number of limitations. The `Target' technique has always been associated with an additive planting design as it involves surrounding a plant of interest (the `target') with other plants (the `neighbours') while including above- and below-ground partitions to prevent root and shoot competition. Most studies using this technique have not provided adequate control for apparatus effects, yet this method appears to have the most potential for application in ecological studies, especially in the field. A standard protocol for the target technique is proposed allowing greater control over potential apparatus effects than in previous studies and allowing assessment of the interaction between root competition and shoot competition.     

Responses of Fraxinus excelsior seedlings to grass-induced above- and below-ground competition

The competitive interactions between woody seedlings and herbaceous vegetation have received increasing interest in recent years. However, little is known about the relative contributions and underlying mechanisms of above- and below-ground competition between species. We used a novel experimental approach to assess the responses of Fraxinus excelsior seedlings to different combinations of root and shoot competition imposed by the grass Dactylis glomerata under greenhouse conditions. Seedling growth was significantly reduced by competition for soil resources, but neither biomass nor height were significantly affected by shoot competition for light. Competitive response indices based on biomass confirmed that below-ground competition was more important than above-ground competition, and indicated that root and shoot competition did not interact to influence plant growth. Fraxinus biomass allocation and seedling traits were almost all significantly affected by root competition; these responses varied depending on the trait examined. In contrast, morphological responses to shoot competition were limited. In the absence of root competition, seedlings showed a significant increase in the biomass allocated to leaves and a greater leaf area ratio in response to shoot competition. Our findings suggest that morphological modifications help to mitigate the negative effects of competition, but the expression of plasticity may be suboptimal due to resource constraints. The present study also highlights the importance of appropriate experimental controls and analysis to avoid confounding effects of experimental design and ontogeny on the interpretation of competitive responses.     

植物种群表观竞争机制及其研究进展

植物表观竞争既影响群落结构和组成,也与生态系统功能密切相关。目前,国内尚缺乏对植物表观竞争的实验研究和机制探索。对植物表观竞争概念和研究进展进行了系统梳理和阶段性总结,分析了草食动物为媒介的表观竞争主要通过食物和生境介导植物竞争过程,以及ELP-采食者关系对表观竞争的影响;植物参与的表观竞争通常与资源竞争共同作用;AM真菌为媒介的表观竞争不仅受到宿主植物菌根依赖性的影响,还与AM真菌对资源的需求和环境资源供给状况有密切关系。未来研究既要注重实验设计的合理性,也要考虑测定方法和评价参数的有效性;重视植物群落和生态系统水平上表观竞争的功能研究。这不仅能够为植物群落结构和物种共存提供机制性解释,而且能够丰富经典的植物竞争理论和多营养级相互作用的食物网理论。     

施肥和放牧对青藏高原高寒草甸物种丰富度的影响

在草地生态系统中, 施肥通常会导致生物多样性下降, 但是关于引起生物多样性下降的机制还存在着很大的争议。该研究基于一个4年的施肥实验, 试图通过个体大小的不整齐性和单位植物氮含量, 定性地揭示青藏高原东部高寒草甸施肥后多样性下降的原因。研究显示: 在封育地, 施肥致使个体大小不整齐性增加了15%, 并不同程度地增加了物种的高度。同时, 施肥使物种间单位植物氮含量存在显著差异的数目降低了65%。施肥后光竞争加剧, 导致大个体植物排斥小个体植物, 进而引起了物种丰富度下降29.6%。与封育地不同, 放牧地施肥并没有改变个体大小不整齐性和物种的高度, 而是使物种间单位植物氮含量存在显著差异的数目增加了11.4%。施肥并没有改变放牧地的光竞争强度, 而是增加了物种间对土壤营养元素氮的竞争强度, 进而引起了物种丰富度下降17.3%。该研究还发现, 放牧施肥地的物种丰富度下降速度和等级显著低于封育施肥地的物种丰富度下降速度和等级, 这表明放牧减缓了施肥对物种丰富度的影响力。     

Comparison of pitfall traps and litter bags for sampling ground-dwelling arthropods

Abstract:  For their simplicity and effectiveness, pitfall traps have become a standard technique to measure the activity and relative abundance of ground-dwelling arthropods. Permeable screen or mesh bags filled with plant material, referred to as litter bags, have also recently been employed as a complementary sampling technique for epigeal taxa. The anticipated need for increased field research on arthropod populations, particularly in transgenic crops with insecticidal properties, suggests that a relative assessment of both sampling methods could contribute to the design of future studies. Comparisons among pitfall traps, and litter bags placed above- or below-ground indicated that above-ground litter bags most frequently succeeded in collecting certain groups of arthropods associated with moisture and sheltered areas, including centipedes (Chilopoda) and beetle larvae (Carabidae, Staphylinidae). Conversely, pitfall traps most often captured taxa considered active at ground level, such as adult carabids or harvestmen (Opiliones). For taxa collected in >40% of all three trap types, bootstrap confidence intervals for the coefficient of variation (CV; used to assess precision or sampling efficiency) showed that above-ground litter bags were significantly more precise than pitfall traps for sampling elongate springtails (Collembola) and adult rove beetles (Staphylinidae), but only during the first year of sampling. While below-ground litter bags often appeared similar to one or both of the other trap types, in no case were below-ground litter bags best based on frequency of collection or CV. Though differences were not always consistent between years, results suggest that the additional effort required to sample using litter bags may be justified for the collection of some ground-dwelling taxa.     

Effect of soil nutrient heterogeneity on the symmetry of belowground competition

Size variability in plant populations has been extensively studied andmuch of this inquiry has focused on the types of competition that are involvedin increasing or decreasing size inequality. It is often assumed thatcompetition for light is size-asymmetric, meaning that a plant canpotentially dominate a competitive relationship through shading if it is tallerthan its competitor. The light resources obtained by the taller plant are thusdisproportionate to its size. In contrast, competition for soil resources maybemore size-symmetric, the amount of soil nutrients obtained seems to be indirect proportion to a plant's size. Most studies examining belowgroundcompetition have used homogeneouslydistributed nutrient resources. Soil homogeneity could makesize-asymmetric belowground competition unlikely, but homogeneity is notoften found in nature. In this study I use a greenhouse pot experimentutilizingIpomoea tricolor to examine the hypothesis thatsize-asymmetric competition for soil nutrients may result when soilresource distribution is spatially heterogeneous. The results did not supportthe hypothesis of belowgroundsize-asymmetric competition. Differences between experimental treatmentsand controls were not statistically significant suggesting size-symmetriccompetition. Implications of these results are discussed.     

植物竞争研究综述

概述了竞争的概念、理论 (包括最大生长率理论、最小资源需求理论和资源动态两阶段说 )、特性 (主要有竞争不对称性和环境相关性 )及其影响因素 ,竞争与群落组成、结构和动态的关系。并对近几十年来生态学上关于植物竞争的争论焦点作了论述 ,即竞争对个体生长、繁殖和存活及群落演替的相对重要性 ,主要介绍了Grime、Newman&Tilman和Deborah&Ariel的 3种观点 ,其中Grime认为竞争不重要 ,相反Newman&Tilman认为竞争比较重要 ,Deborah&Ariel则提出了资源动态两阶段说 ,认为竞争重要与否取决于两个条件 :一为植物对土壤资源的吸收利用与非生物因素两者分别在多大程度上决定了资源可利用性的渐次降低 ;二为资源正波动期因竞争引起的生长量减少在多大程度上决定了资源负波动期个体的存活率     

塔里木荒漠优势植物——胡杨种内、种间竞争研究

采用逐步扩大范围的方法确定影响对象木的邻体最佳竞争范围,利用Hegyi提出的单木竞争指数模型对塔里木荒漠优势植物———胡杨种内、种间竞争强度进行定量分析。结果表明,在胡杨群落中,对象木的最适竞争距离为6 m,胡杨种内、种间竞争强度随着植株径级的增大而逐渐减小,种内竞争较种间竞争剧烈。竞争强度与对象木胸径服从幂函数关系,从模型预测结果可知当胡杨胸径达30 cm以上时,竞争强度变化很小。因此,应在此前采取适当的人工疏伐措施来促进植株生长和增强生态系统的稳定性。