植物群落物种共存机制的研究进展

物种共存是由进化、历史及生态尺度上的过程决定的.现存的理论从不同的尺度探讨了植物群落物种共存的可能机制,本文阐述了其中几种重要的理论.种库理论在进化和历史尺度上解释了植物物种共存的形成原因.在生态尺度上,虽然传统的生态位理论受到质疑,但是更新生态位理论和资源比率/异质性假说越来越受到重视;竞争共存理论认为具有相似竞争能力或能够避免竞争排斥的植物物种可以共存.在非平衡条件下,生物和非生物因素对植物物种共存也有显著影响,它们一方面作用于竞争优势种,使竞争优势向稀有种转移,另一方面可以创造生境时空异质性,为生态位分化提供机会.生态漂变学说认为群落中物种的组成不断变化,物种的共存和分布由随机因素决定.这些植物群落物种共存理论各有所长,互相补充.应用现代科学技术进行研究,结果必将促进人们对植物物种共存问题的深刻理解.     

土壤水分变化对黄土丘陵区撂荒群落共存种竞争的影响

群落共存种对限制性资源的竞争响应是群落组建、维持与演替的重要影响因素。以黄土丘陵区摞荒群落共存种为研究对象,研究不同水分条件下的竞争响应差异,以期揭示生境变化时群落演替的植物竞争机理。试验设土壤水分与混植密度两个处理,其中混植密度采用CDS （Community density series）法,含3个密度水平（衡水试验为3、12、21株/m²,变水试验为5、20、35株/m²）;土壤水分设衡水（高（75% FC）、中（60% FC）、低（45% FC）3个水平）与变水（由高水（75% FC）经50 d持续干旱至低水（45% FC））两种处理。根据共存种株高的阶段性生长与其在群落中的相对生物量等生长表现,在CDS法基础上扩展并测算了生长变化量、相对生物量和相对生长率等指标。据此,研究竞争效应和水分胁迫效应的相对影响,共存种相对竞争能力和竞争等级对土壤水分变化的响应,及高水期因竞争引起的个体大小不对称性与低水期死亡率的关系。结果表明：1）不同水分多种混植共存条件下,竞争效应对多数共存种在群落中的生长表现较水分胁迫效应影响更大;以相对生物量为指标的竞争的群落效应（4.30±0.49）较水分胁迫的群落效应（-23.56±20.38）大,且二者累加效应（4.76±0.4）大于二者单项效应之和（-19.26±20.48）,即竞争+水分胁迫对多种共存群落具有协同效应。2）当水分条件发生变化时,群落共存种的竞争等级并不完全相同（P > 0.05）,相应的会发生变化,且水分差别越大,共存种竞争等级的差别也越大;高水条件下群落共存种竞争所引起的群落效应最大（4.84±0.23）,部分支持最大生长率理论,即限制性资源水平越高,群落共存种竞争越激烈。3）变水试验中,10种群落共存种在高水期因竞争引起的个体大小差异与低水期的死亡率呈正相关关系（P=0.015,r²=0.801）,根据资源动态两阶段假说,说明试验条件下群落共存种竞争是群落组建的重要因素。     

群落中克隆植物的重要性

综述了群落水平上克隆植物的重要性,克隆植物的生态习性,克隆植物的竞争关系等方面的研究进展,并试图在克隆植物竞争关系的背景下,结合其生境状况,探讨植物克隆性与植物物种多样性的关系。克隆植物的等级系统（基株—分株系统—分株）使其具有克隆性所赋予的多样的生活史、资源利用以及空间占据方式。列举了克隆植物的特性及其在群落中的作用。这些例子表明,克隆性强烈地影响和制约着植物群落的空间格局与竞争关系。然而克隆性也能通过权衡活动性与局部持久性来缓解对物种共存的抑制。克隆植物具有在时空尺度上分株间相互调节的机制,直接体现在群落中小尺度（个体与个体间）与大尺度（种群与种群间）间的相互作用上。丰富了传统的竞争和生态位划分理论,为群落中物种共存提供了合理解释。因此克隆植物在群落中的出现拓宽了群落系统潜在机制的范围。     

竞争-耐胁迫-杂草型植物对策理论及其应用研究进展

竞争-耐胁迫-杂草型植物(CSR)对策理论主要研究植物对环境胁迫和干扰的适应性特化。从功能生态学角度讲,CSR理论是通过解释各种功能性状协同变异的主导维度来反映植物的生态对策。而物种所采取的生态对策将决定其在群落中的生态位,并最终会影响生态系统过程与功能。自1974年Grime首次提出CSR对策的概念以来,CSR对策思想就受到了生态学家很大的关注,经过近半个世纪的发展,目前已经形成了比较完整的理论和研究方法体系。试图在回顾CSR理论及其研究方法发展的基础上,阐述其基本内涵,并重点从功能性状变异空间、植物适应性、群落过程和生态系统特性等4个方面概述CSR理论及其分析模型的验证及应用,梳理其最新研究进展。CSR理论已经得到了很多研究的广泛验证并应用于多种尺度上的植物生态学研究,但仍有一些方面值得进一步发展完善,如第四区缺失问题、功能性状变异主轴以外的变异、CSR对策与功能性状种内变异和谱系关系的联系等方面需要深入研究。     

黄土丘陵区不同立地条件下猪毛蒿种内、种间竞争

猪毛蒿(Artemisia scoparia)通常是黄土丘陵区撂荒演替前期群落优势种,在无人为干扰的情况下,猪毛蒿群落通常会向冰草(Agropyron cristatum)群落或阿尔泰狗娃花(Heteropappus altaicus)群落,或长芒草(Stipa bungeana)群落等演替。该文通过河阶地和梁峁阴坡中猪毛蒿生长特征的调查和种内、种间竞争田间试验,从植物竞争角度对猪毛蒿群落的演替机制给予了解释。结果表明:1)两类样地中猪毛蒿的生长都趋于小型化,有少数大个体和多数小个体,都存在异速生长现象,说明两类样地存在竞争,且对猪毛蒿的生长形态具有塑造作用; 2)由于单位地上生物量竞争效应排除了立地条件和个体大小的影响,因而比总竞争效应更能说明种对间的相对竞争能力。梁峁阴坡地和一、二级河阶地三种立地条件下,7种测试植物中对猪毛蒿的相对竞争能力以演替后期多年生植物较高,而演替前期一年生植物较低,说明演替后期种对前期种的竞争抑制是演替驱动力之一; 3)以各测试植物对猪毛蒿单位重量竞争抑制程度平均值来看,以梁峁阴坡地最大,二级河阶地次之,一级河阶地最小,说明立地条件越差,土壤资源可利用水平越低,竞争越激烈; 4)一级河阶地和梁峁阴坡地各测试植物对猪毛蒿的竞争等级发生了显著变化,说明环境条件差别较大时,植物的竞争等级会发生变化。     

植物竞争研究进展

竞争系指两个以上有机体或物种间阻碍或制约的相互关系。它是塑造植物形态、生活史的主要动力之一;并对植物群落的结构和动态具有深刻的影响。因其在生态学的中心地位,生态学家已从不同的侧面研究了这一复杂的生态学现象;生态学也因此而得到了发展。然而,人们对竞争的理解不尽一致,因而导致了概念上的混乱,平行研究相对缺乏、不同研究间的比较困难,从而阻碍了学科的发展。本文试就植物竞争的概念、竞争理论、竞争研究的实验方法、影响竞争能力的主要因素、种内和种间竞争对种群和群落的影响,如竞争与物种共存等进行综述;我们在总结研究成就的同时,亦指出了现有研究的局限性。     

植物竞争研究进展

竞争系指两个以上有机体或物种间阻碍或制约的相互关系。它是塑造植物形态、生活史的主要动力之一;并对植物群落的结构和动态具有深刻的影响。因其在生态学的中心地位,生态学家已从不同的侧面研究了这一复杂的生态学现象;生态学也因此而得到了发展。然而,人们对竞争的理解不尽一致,因而导致了概念上的混乱,平行研究相对缺乏、不同研究间的比较困难,从而阻碍了学科的发展。本文试就植物竞争的概念、竞争理论、竞争研究的实验方法、影响竞争能力的主要因素、种内和种间竞争对种群和群落的影响,如竞争与物种共存等进行综述;我们在总结研究成就的同时,亦指出了现有研究的局限性。     

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。     

半湿润常绿阔叶林次生演替阶段植物多样性和群落结构特征

以飒马场不同次生演潜阶段的半湿润常绿阔叶林为对象,测定物种丰富度、群落数量特征指标,分析植物多样性与群落结构特征之间的关系.结果表明,随着植物群落演替,物种丰富度不断增加,物种丰富度最高演替阶段的乔木层物种达到24个;植物多样性与植物个体密度间呈正相关关系;乔木层植物个体平均高与植物多样性间呈负相关关系;群落郁闭度和胸高断面积与植物多样性之间则呈对数函数关系.植物多样性与优势种个体密度和群落地上部分现存量间未呈现一致性规律.植物种-个体数间呈幂函数关系.随着植物多样性增加,种间竞争加剧.每个植物种为了维持临界最低种群,可能采取最大种群策略,导致群落植物个体数增加,密度增大,从而改变了群落结构特征.     

植物病原菌在森林动态中的作用

植物病原菌作为森林生态系统的重要组成成分及调控因子之一,在森林动态中扮演着重要的角色。植物病原菌通过侵染过程导致寄主植物的幼苗及成熟个体死亡、成熟个体的种子量降低或不实,或造成植物个体或群落中不同物种不同程度的病害,影响它们之间的营养竞争,从而导致群落结构、物种及个体数量的变化。感染散布前、后的种子和土壤种子库中的种子,以及由种子萌发产生的幼苗,它们的存活率降低,进而影响森林中的种子散布、幼苗更新与增补格局。在天然林中,先锋树种比顶极树种对病原菌更敏感,群落演替的早期阶段对病原菌比较敏感。植物病原菌主要通过密度依赖机制造成森林树种不同的死亡格局,从而参与森林的动态过程。     

Intensity and Importance of Competition for a Grass (Festuca rubra) and a Legume (Trifolium pratense) Vary with Environmental Changes

How plant competition varies across environmental gradients has been a long debate among ecologists. We conducted a growth chamber experiment to determine the intensity and importance of competition for plants grown in changed environmental conditions. Festuca rubra and Trifolium pratense were grown in monoculturs and in two- and/or three-species mixtures under three environmental treatments. The measured competitive variations in terms of growth (height and biomass) were species-dependent. Competition intensity for Festuca increased with decreased productivity, whilst competition importance displayed a humpback response. However, significant response was detected in neither competition intensity nor importance for Trifolium. Intensity and importance of competition followed different response patterns, suggesting that they may not be correlated along an environmental gradient. The biological and physiological variables of plants play an important role to determine the interspecific competition associated with competition intensity and importance. However, the competitive feature can be modified by multiple environmental changes which may increase or hinder how competitive a plant is.     

Competition components along productivity gradients – revisiting a classic dispute in ecology

Competition is ubiquitous in plant communities with various effects on plant fitness and community structure. A long-standing debate about different approaches to explain competition is the controversy between David Tilman and Philip Grime. Grime stated that the importance of competition relative to the impact of the environment increases along a productivity gradient, while Tilman argued that the intensity of competition is independent of productivity. To revisit this controversy, we assumed that the effects of plant–plant interactions are additive and applied the new competition indices by Díaz-Sierra et al. (2017) in a field experiment along a productivity gradient in S-Germany, using the rare arable plant Arnoseris minima as a study species. The ‘target technique' was applied, to separate the effects of root and shoot competition. The study plants were exposed to five competition treatments with three replicates in 18 sites, respectively. We investigated the expectation that root competition is more intense in unproductive sites than shoot competition. Additionally, we predicted survival to be less affected by competition than growth-related plant parameters. Using the biomass of individuals without competition as a proxy for site productivity there was a positive relationship with competition importance but no relationship with competition intensity when plants experienced full competition. Survival of the target plants was unaffected by competition. Root competition was the main mechanism determining the performance of the target plants, whereas the effect of shoot competition was relatively low albeit increasing with productivity. We conclude that when considering plant–plant interactions additive both Grime's and Tilman's theories can be supported.     

The shifting balance of facilitation and competition affects the outcome of intra- and interspecific interactions over the life history of California grassland annuals

Trait-based resource competition in plants, wherein more similar plants compete more strongly for resources, is a foundation of niche-based explanations for the maintenance of diversity in plant communities. Alternatively, neutral theory predicts that community diversity can be maintained despite equivalent resource requirements among species. We examined interactions at three life history stages (germination, survival, and juvenile-adult growth) for three native and three exotic California annual species in a glasshouse experiment. We varied plant density and species composition in small pots, with pots planted with either intraspecific seeds or in a three species mix of intra- and interspecific neighbors. We saw a range of facilitative, neutral, and competitive interactions that varied significantly by species, rather than by native or exotic status. There were more competitive interactions at the emergence and juvenile-adult growth stages and more facilitative interactions for survival. Consequently, the relative strength of competition in intraspecific versus mixed-species communities depended on whether we considered only the juvenile-adult growth stage or the entire life history of the interacting plants. Using traditional analysis of juvenile-adult growth only, all species showed negative density-dependent interactions for final biomass production. However, when the net effect of plant interactions from seed to adult was considered, which is a prediction of population growth, two native species ceased to show negative density dependence, and the difference between intraspecific and mixed-species competition was only significant for one exotic species. Results were consistent with predictions of neutral, rather than niche, theory for five of six species.     

Counterbalancing effects of competition for resources and facilitation against grazing in alpine snowbed communities

Alpine snowbeds are habitats where the major limiting factors for plant growth are herbivory and a small time window for growth due to late snowmelt. Despite these limitations, snowbed vegetation usually forms a dense carpet of palatable plants due to favourable abiotic conditions for plant growth within the short growing season. These environmental characteristics make snowbeds particularly interesting to study the interplay of facilitation and competition. We hypothesised an interplay between resource competition and facilitation against herbivory. Further, we investigated whether these predicted neighbour effects were species‐specific and/or dependent on ontogeny, and whether the balance of positive and negative plant–plant interactions shifted along a snowmelt gradient. We determined the neighbour effects by means of neighbour removal experiments along the snowmelt gradient, and linear mixed model analyses. The results showed that the effects of neighbour removal were weak but generally consistent among species and snowmelt dates, and depended on whether biomass production or survival was considered. Higher total biomass and increased fruiting in removal plots indicated that plants competed for nutrients, water, and light, thereby supporting the hypothesis of prevailing competition for resources in snowbeds. However, the presence of neighbours reduced herbivory and thereby also facilitated survival. For plant growth the facilitative effects against herbivores in snowbeds counterbalanced competition for resources, leading to a weak negative net effect. Overall the neighbour effects were not species‐specific and did not change with snowmelt date. Our finding of counterbalancing effects of competition and facilitation within a plant community is of special theoretical value for species distribution models and can explain the success of models that give primary importance to abiotic factors and tend to overlook interrelations between biotic and abiotic effects on plants.     

太白红杉种内和种间竞争研究

采用逐步扩大范围的方法确定影响对象木 (Objectivetree) 的最佳竞争范围, 利用单木竞争指数的改进模型对太白红杉 (Larixchinensis) 种内和种间竞争强度进行了定量分析, 并讨论了不同竞争强度下太白红杉的形态变化。结果表明 :随对象木胸径的增大, 由于太白红杉种群自然稀疏过程中密度调节作用, 植株距离增加, 种内竞争强度降低 ;太白红杉主要分布于亚高山地段, 群落内其它物种较少, 个体普遍较小, 结果种间竞争相对较弱, 种内与种间竞争关系顺序为 :太白红杉 太白红杉 >巴山冷杉 (Abiesfargesii) 太白红杉 >牛皮桦 (Betulaplatyphylla) 太白红杉 >其它树种 太白红杉 ;竞争强度和对象木胸径的关系服从幂函数关系 (CI =AD-B), 当太白红杉胸径达到 35cm以上时, 竞争强度几乎没有变化, 所得的预测模型能很好地预测太白红杉种内和种间的竞争强度 ;不同竞争强度下, 太白红杉主茎各层的分枝角度、总分枝数、当年生枝条长、平均枝长和活枝数均表现出显著的差异。表明采用逐步扩大范围的方法能有效地确定竞争木范围, 较好地反应太白红杉种内和种间的竞争关系。同时, 太白红杉通过自身形态变化, 提高了对光的截获能力和对不同竞争强度的适应能力。     

Experimental evidence for root competition effects on community evenness in one of two phytometer species

Aims Plant–plant interactions, being positive or negative, are recognized to be key factors in structuring plant communities. However, it is thought that root competition may be less important than shoot competition due to greater size symmetry belowground. Because direct experimental tests on the importance of root competition are scarce, we aim at elucidating whether root competition may have direct or indirect effects on community structure. Indirect effects may occur by altering the overall size asymmetry of competition through root–shoot competitive interactions.     

Linking individual response to biotic interactions with community structure: a trait-based framework

1.  Due to species-specificity of the outcomes of biotic interactions, it is difficult to generalize from observed biotic interactions at the individual plant level to the effect of those interactions at the community level. To evaluate the importance of biotic interactions in shaping plant communities, it is necessary to understand how the outcomes of the complex interactions observed at the individual level can influence community structure.
2.  Here, we propose a trait-based framework that identifies and organises mechanisms affecting community structure (here described with relative abundances of plant functional traits – i.e. the distribution of trait values at the community level). We applied our approach to a single leaf trait, specific leaf area (SLA), to link individual responses to plant interactions with community structure (SLA distribution observed at the community level) and to test whether biotic interactions can predict the functional composition of subalpine grasslands. We evaluated the generality of our model through a cross-validation with a set of eight subalpine grasslands independent from the four fields used to build the model.
3.  We found that competition and facilitation were able to explain the functional composition of subalpine grasslands, and the relevant fitness components (survival or growth) explaining this link changed depending on the limiting resources. When soil water availability was limiting, positive plant-plant interactions acting on survival were able to explain community structure. In contrast, when no water limitation was observed competition acting on individual growth was the main driver of community structure.
4.  Our framework enables evaluation of the consequences of biotic interactions observed at individual level on community structure, thereby indicating when and where different types of plant-plant interactions are important.     

Temperature‐dependence of minimum resource requirements alters competitive hierarchies in phytoplankton

Resource competition theory is a conceptual framework that provides mechanistic insights into competition and community assembly of species with different resource requirements. However, there has been little exploration of how resource requirements depend on other environmental factors, including temperature. Changes in resource requirements as influenced by environmental temperature would imply that climate warming can alter the outcomes of competition and community assembly. We experimentally demonstrate that environmental temperature alters the minimum light and nitrogen requirements – as well as other growth parameters – of six widespread phytoplankton species from distinct taxonomic groups. We found that species require the most nitrogen at the highest temperatures while light requirements tend to be lowest at intermediate temperatures, although there are substantial interspecific differences in the exact shape of this relationship. We also experimentally parameterize two competition models, which we use to illustrate how temperature, through its effects on species’ traits, alters competitive hierarchies in multispecies assemblages, determining community dynamics. Developing a mechanistic understanding of how temperature influences the ability to compete for limiting resources is a critical step towards improving forecasts of community dynamics under climate warming.     

AM真菌在草原生态系统中的功能

AM真菌是土壤生态系统中重要的微生物类群,能与陆地生态系统中80%以上的高等植物建立共生体系。目前,AM真菌在维持草原生态系统稳定性中的功能已经成为生态学研究的热点问题之一。基于此,从植物个体、种群、群落和生态系统等不同层次探究AM真菌在维持植物群落多样性和草原生态系统稳定性中的功能。分析发现在个体水平上,AM真菌对宿主植物具有促生效应、抑制效应或中性效应。在种群水平上,分析AM真菌对不同宿主植物吸收土壤矿质营养的分配和调控策略,围绕构成草原植被的两大组成成分:牧草和有毒植物,论述AM真菌对植物种群增长和衰败的调控机制,并从草原植物群落的物种多样性和稳定性角度,探讨AM真菌与植物群落之间的相关性。在生态系统水平上,围绕AM真菌对草原生态系统的演替和退化草原的修复等展开论述,以期为利用AM真菌开展草原生态系统保护和恢复治理提供理论依据,并对草原菌根生态学领域未来的研究进行展望。