Inference from plant competition experiments: the effect of spatial covariance

Two-species plant competition experiments are often drawn on in an attempt to understand and possibly to predict the formation of plant community structures. Frequently, such competition experiments use a non-random spatial design but is analysed in models where a random spatial design is implicitly assumed. Here, the effect of spatial covariance in plant competition experiments on the inferred ecological conclusions is demonstrated. It is shown that spatial covariance, which is dependent on the competition kernel and the experimental design, may affect the estimation of parameters of interest. Based on the same set of experimental data, the predicted outcome of competition between two Arabidopsis thaliana genotypes can vary between (almost certain) coexistence and (almost certain) exclusion of one genotype, depending on the assumptions made about the spatial scale of competition between the plants . The predicted outcome of competition depended strongly on the mean interaction distance of the competition kernel, whereas the functional shape of the competition kernel was less important. It is suggested that plants in manipulated competition experiments should be placed randomly, such that the design of the competition experiment is in agreement with the model used in the analysis.     

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.     

昆虫种间表观竞争研究进展

表观竞争是资源竞争以外的一种新型的种间关系,是指由共同享有的自然天敌中介的、物种之间在种群数量上表现出明显负效应的现象。物种之间的表观竞争被认为和资源竞争具有同等的重要性,尤其是对植食性昆虫来说更为重要。近十年来关于昆虫表观竞争的理论和实验研究进展迅速,已成为昆虫生态学研究中的一个新的生长点。本文从表观竞争的定义、实验研究实例、理论模型、作用机制及共存机制等方面,对近些年来关于昆虫表观竞争研究的进展进行了概括和总结,并对表观竞争理论在害虫防治中的应用前景进行了展望。     

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。     

基于引力模型的林木竞争分析

引力模型是否可以应用于森林群落林木竞争关系分析是值得研究的问题。基于引力模型建立林木相对活力圈能反映竞争木活力大小,基于竞争木的相对活力圈建立引力竞争指数能准确反映林木生长与林木竞争的关系。以浙江省天目山国家级自然保护区针阔混交林为研究对象,将V_Hegyi竞争指数、引力竞争指数分别与胸径进行相关分析,胸高断面积生长量分别与2种竞争指数进行相关分析,胸径生长率与2期引力竞争指数的比值(2021年与2006年的引力竞争指数之比)进行相关分析,此外,对相对活力圈直径与胸径进行相关分析,并比较分析了活立木与枯死木的竞争指数大小。结果表明: 2种竞争指数与胸径均呈显著负相关,且均服从幂函数关系。林木胸高断面积生长量与2种竞争指数均呈显著负相关,但引力竞争指数比V_Hegyi竞争指数更能反映林木生长与林木竞争的关系。相对于V_Hegyi竞争指数的比值,2期引力竞争指数的比值更能说明林木生长与林木竞争的关系。在针阔混交林中,阔叶树种的生长与竞争的相关性>针叶树种生长与竞争的相关性。林木枯损受竞争的显著影响。林木相对活力圈大小与林木胸径大小呈显著负相关。引力模型是反映空间相互作用的重要模型之一,可以应用于林木竞争关系的研究,且基于引力模型建立的引力竞争指数可以作为评价林木竞争和林木活力的一个空间结构指标,比V_Hegyi竞争指数更能反映林木生长与林木竞争的关系。     

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。     

种间竞争的一个新的数学模型—对经典的Lotka—Vol—terra竞争方程的扩充

本文根据营养动力学理论,建立了一类种间竞争的新的数学模型:它是单种群增长的Cui-Lawson模型,在种间竞争上的推广。新的种间竞争模型克服了经典的种间竞争的Lotka-Volteira方程的局限与不足,具有更广泛和复杂的行为,并在特殊条件下以Lotka-Volterra竞争方程为其特例。因此,新的种间竞争的数学模型是更一般的解释性模型,是对经典的Lotka-Voterra竞争方程的扩充。     

The evolution of sperm-allocation strategies and the degree of sperm competition

The prevailing viewpoint in the study of sperm competition is that male sperm-allocation strategies evolve in response to the degree of sperm competition an ejaculate can expect to experience within a given mating. If males cannot assess the degree of sperm competition their ejaculate will face and/or they are unable to facultatively adjust sperm investment in response to perceived levels of competition, high sperm allocation (per mating) is predicted to evolve in the context of high sperm competition. An implicit assumption of the framework used to derive this result is that the degree of sperm competition is unaffected by changes in sperm-allocation strategies. We present theory based on an alternative perspective, in which the degree of sperm competition and the sperm-allocation strategy are coupled traits that coevolve together. Our rationale is that the pattern of sperm allocation in the population will, in part, determine the level of sperm competition by affecting the number of ejaculates per female in the population. In this setting, evolution in sperm-allocation strategies is driven by changes in underlying environmental parameters that influence both the degree of sperm competition and sperm allocation. This change in perspective leads to predictions that are qualitatively different from those of previous theory.     

黄山松种群邻体范围与邻体竞争强度的研究

当环境中可利用的资源低于种群最佳生长需要时,种群内个体之间就会产生竞争。竞争常发生在邻近植物之间,植物地上部分和地下部分的竞争范围是不同的。目前,有很多度量竞争强度和确定竞争范围的方法,能否对这些方法进行改进,提出更具有说服力的方法,是本研究的出发点。黄山松是常见的针叶树种,分布范围广,生存能力强,是亚热带中部中山地区代表群系的建群种,也是较高海拔地区重要的造林树种,具有较强的代表性。对黄山松的邻体竞争研究发现:(1)改进的竞争指数有更强的说服力;(2)逐步扩大范围的方法能很好地确定植株间的竞争范围;(3)邻体竞争强度随影响范围的增加而增加,在一定的范围内增加的较快,而超出该范围后增加的幅度变小,可以此为依据来确定邻体范围;(4)不同径级的基株,邻体范围有一定的差异;(5)邻体竞争强度和影响范围的关系服从对数函数关系(CI=AlnC+B)。结果表明,改进的竞争指数能很好地度量竞争强度,采用逐步扩大范围的方法能有效地确定邻体范围。     

Individual-Based Competition Between Species with Spatial Correlation and Aggregation

In order to clarify the theoretical relationship between individual behavior and population-level competition between two species with spatial correlation, this paper describes how discrete-time competition equations for the two species can be derived from local resource competition among individuals. Competition type of each species is either scramble, contest, or modified contest, and for various combinations of two competition types, different competition models are derived. Simple competition models that can approximate the above models when competition is weak are also derived. Furthermore, the derived models are used to investigate how coexistence conditions and coexistence probability depend on spatial correlation and aggregation of individuals. For the weak competition models, spatial aggregation and non-correlation, in terms of measures adopted here, play exactly symmetric roles in promoting coexistence. In contrast, for the fully developed models, spatial aggregation generally exerts stronger effects than non-correlation on coexistence. Coexistence probability also depends greatly on competition types. For example, two species are generally more likely to coexist when they are of the same competition type than of different competition types. Coexistence probabilities from the mathematical analysis are in good agreement with those from individual-based simulations.     

A review of competition in north temperate dung beetle communities

Abstract. 1. Studies of north temperate dung beetle communities frequently invoke competition as an influential ecological process. In this review, the evidence for competition in north temperate dung beetle communities is evaluated and the role of competition as a factor affecting community structure is assessed.
2. Resource limitation and the evidence for interspecific competition are assessed by collating the available experimental and observational evidence for both the adult and larval stages of the dung beetle life cycle. The role of competition as a structuring force in dung beetle communities is discussed under the following headings: niche dynamics, migration to and from individual pats, the aggregation model of co-existence, and metapopulations.
3. Some of the main conclusions are that competition for space is much more likely to occur than competition for food; the effects of competition on community structure are poorly understood; several of the influential studies of competition in north temperate dung beetle communities need to be evaluated carefully. The differences in ecology between tropical and temperate dung beetle communities are clarified.
4. As priorities for future research, resource utilisation and competition should be researched experimentally: density-dependent relationships should be investigated, particularly for the larval stages, as should competitive interactions with other dung fauna. If such experimental approaches establish convincingly the occurrence of competition, then the extent of competition in the field and under real world conditions needs to be established. A functional group classification of dung beetles and other dung fauna is described, which may improve the generality of interpretation from individual, site-specific results.     

The intensity of competition versus its importance: an overlooked distinction and some implications

The intensity of competition is a physiological concept, related directly to the well-being of individual organisms but only indirectly and conditionally to their fitness, and even more indirectly to the evolution of populations and the structure of communities. The importance of competition is primarily an ecological and evolutionary concept, related directly to the ecology and fitness of individuals but only indirectly to their physiological states. The intensity of competition is not necessarily correlated with the intensities of predation, disturbance, abiotic stress, or other ecological processes. The importance of competition is necessarily relative to the importances of other processes. Intensity refers primarily to the process of present competition, whereas importance refers primarily to the products of past competition. The distinction between the intensity and the importance of competition clarifies two long-standing ecological debates. Some ecologists have proposed that competition is greater in more stressful habitats, others the opposite, and still others that no such relationship exists. Evidence cited to refute or support these positions often confuses intensity and importance. Distinguishing between them focuses questions more sharply and indicates what sorts of new evidence should be sought. The more widely known debate over the prevalence of competition as an agent of community structure is a debate about the importance of competition, but evidence about the intensity of competition has often been used by both sides. We argue that intensity and importance need not be correlated, and so measurements of the intensity of competition are not directly relevant to this debate. This distinction also generates testable hypotheses and suggests directions for research. For example, we hypothesize that competition can be unimportant even if it is very intense: no such hypothesis is possible unless importance is distinguished from intensity. We discuss the application of these ideas to methods and theories used to study competition, ecological communities, and the evolution of competitive ability. We advocate a research approach that presumes multiple, interacting causes, including competition, affecting community structure, and we show how the distinction between intensity and importance helps to make this feasible.     

基于竞争和扩散能力的非捕食-被捕食集合种群系统的竞争机制

对于非捕食 被捕食(食饵)生态系统,强弱物种之间存在一定的竞争影响.在不考虑栖息地毁坏的情况下,引进双向竞争机制,将Tilman的单向竞争模式推广为n集合种群双向竞争模型,并对6-集合种群的竞争动态进行了计算机模拟研究.结果表明,在平衡态,种群竞争共存的条件是其竞争能力与扩散能力呈现指数型负相关关系,竞争的结果使物种的强弱序列发生变化;物种竞争排除与共存受迁移扩散能力和竞争能力影响很大,在局域斑块上竞争排斥的集合种群在广域尺度上可以竞争共存,即逃亡共存.     

Increased competition does not lead to increased phylogenetic overdispersion in a native grassland

That competition is stronger among closely related species and leads to phylogenetic overdispersion is a common assumption in community ecology. However, tests of this assumption are rare and field‐based experiments lacking. We tested the relationship between competition, the degree of relatedness, and overdispersion among plants experimentally and using a field survey in a native grassland. Relatedness did not affect competition, nor was competition associated with phylogenetic overdispersion. Further, there was only weak evidence for increased overdispersion at spatial scales where plants are likely to compete. These results challenge traditional theory, but are consistent with recent theories regarding the mechanisms of plant competition and its potential effect on phylogenetic structure. We suggest that specific conditions related to the form of competition and trait conservatism must be met for competition to cause phylogenetic overdispersion. Consequently, overdispersion as a result of competition is likely to be rare in natural communities.     

临安区不同森林类型竞争指数比较研究

竞争对树木生长有重要影响,通常用竞争指数定量描述树木之间的竞争关系。以浙江省杭州市临安区的乔木林为对象,利用2004年临安区森林资源监测数据,基于Voronoi图的Hegyi竞争指数,在区域尺度上对临安区多种森林类型的竞争关系进行分析。结果表明:在单木水平上,不同森林类型单木的总竞争指数、种内竞争指数、种间竞争指数在胸径5—10 cm最大;单木平均竞争指数随着胸径的增大而降低,胸径>30 cm的竞争压力得以缓解;临安区林木的竞争主要来源于种内竞争,天然林的种内竞争占总竞争指数的比例>50%,人工林的种内竞争占总竞争指数的比例>60%。径级平均总竞争指数、种内竞争指数、种间竞争指数与胸径的关系均服从对数函数关系。在林分水平上,不同森林类型的林分总竞争指数、种内竞争指数、种间竞争指数存在显著差异。平均总竞争顺序为:天然阔叶林>天然针阔混交林>人工针叶林>天然针叶林>人工针阔混交林>人工阔叶林。种内平均竞争顺序为:人工针叶林>天然阔叶林>天然针叶林>天然针阔混交林>人工针阔混交林>人工阔叶林。种间平均竞争顺序为:天...     

Does experience with competition matter? Effects of source competitive environment on mean and plastic trait expression in Erodium cicutarium

Interspecific competition is likely to act as an agent for selection on local scales, although evidence in plants is sparse so far. We hypothesize that in annual shade-avoiding grassland species, heterogeneity in the intensity of aboveground competition for light may shape patterns of genetic variation and induce phenotypic plasticity in traits affecting competitive ability. We collected maternal seed families of Erodium cicutarium from replicated high and low competition environments and exposed them to different levels of aboveground competition in a glasshouse. We examined effects of seed source and competition treatment on expression of plant traits related to competitive ability and fitness. Source environments with high levels of competition were significantly more heterogeneous in competition intensity at both intermediate (approx. 10 m) and small (approx. 0.1 m) spatial scales. Seed source and competition treatment both had highly significant effects on trait expression. Greater intensity of competition experienced by maternal plants was coupled with lower vegetative biomass production and slower growth rates, and at the same time lower abortion rates in the offspring. We interpret these findings as an indication of greater reproductive efficiency in the next generation, in response to competition experienced by parents. There was higher total phenotypic variability in the plants from high competition source sites, but equivalent levels of phenotypic plasticity across source-site competition levels; no costs of phenotypic plasticity were detected. We concluded that differences in competition intensity can lead to trait differentiation in the next generation. For E. cicutarium, experience with competition matters: it leads to substantial phenotypic differences and more total variability in the offspring generation.     

珍稀濒危植物梓叶槭种群径级结构与种内种间竞争关系

该研究对四川省珍稀濒危植物梓叶槭(Acer catalpifolium)种群的主要分布区进行调查,分析梓叶槭种群径级结构与种内和种间的竞争关系,探讨竞争强度与径级和距离之间的关系。结果表明:(1)梓叶槭胸径与树高之间存在显著的幂函数关系;种群径级结构呈正态分布,中小径级树木较多,高径级树木较少。(2)梓叶槭种内和种间竞争强度分别占总竞争强度(222.87)的15.16%和84.84%,说明竞争主要来自种间。(3)梓叶槭与主要伴生种之间的竞争强度大小依次为柳杉桢楠白栎刺楸灯台树桦木杉木厚朴光皮桦;对象木和竞争木距离与竞争指数之间较好地服从指数函数关系,当对象木与竞争木距离小于1m时,竞争指数可达到5.5,随着距离逐渐增加,其竞争指数相应降低,并最终趋于平缓。(4)竞争强度随对象木胸径的增大而减小,当对象木胸径小于20cm时,受到的竞争压力最大,竞争指数最大的分别是梓叶槭与整个林分(234.98)以及梓叶槭与种间(184.01);当对象木胸径小于10cm时,竞争指数均可以达到18;当对象木胸径大于20cm时,竞争强度变化很小且竞争指数较低;竞争强度与对象木胸径服从幂函数关系(CI=AD-B)。(5)模型预测结果表明,随着梓叶槭胸径的增加,竞争指数均越来越小,竞争强度呈降低趋势;当梓叶槭胸径为0~5cm时,梓叶槭与整个林分受到的竞争强度最大,竞争指数为7.14,占总竞争强度的50%;当胸径大于20cm时,竞争指数变化不大;该模型能很好的预测梓叶槭种内和种间竞争强度。     

Diminishing returns in social evolution: the not-so-tragic commons

A challenge for evolutionary theory is to understand how cooperation can occur in the presence of competition and cheating, a problem known as the tragedy of commons. Here I examine how varying the fitness returns from reproductive competition or cooperation affects the negative impact of competition on a social group. Varying linear returns does not affect the impact of competition. However, diminishing returns, where additional investments in either competition or cooperation give smaller and smaller rewards, reduce the effects of competition on the group. I show that diminishing returns are common in many systems, including social vertebrates, microbes, social insects and mutualisms among species. This suggests that the tragedy of the commons is not so tragic and that the disruptive effects of competition upon social life will often be minor.     

Analysis of species coexistence co-mediated by resource competition and reproductive interference

Reproductive interference is any interspecific sexual interaction that reduces the reproductive success of females through promiscuous reproductive activities of heterospecific individuals. This phenomenon is ubiquitous in nature in both plants and animals, and is frequently observed in biological invasions. However, its effects on interspecific competition remain incompletely understood despite growing concern. To study the interactive effects of resource competition and reproductive interference on species coexistence and exclusion, we analyzed a unified competition model including both processes in symmetric and asymmetric scenarios. The results of our model showed that resource competition and reproductive interference act synergistically to promote competitive exclusion. We also found that when the two processes are asymmetric, the species that is superior in reproductive interference can coexist with or even exclude the species that is superior in resource competition. Therefore, coexistence is possible via an unbalanced trade-off between resource use and reproduction. Our results suggest that integration of reproductive interference and resource competition will contribute to a better understanding of interspecific competition and to more effective biodiversity conservation against management of biological invasions.