代谢互养关系在维持微生物物种多样性中的作用

互养关系（cross-feeding）是微生物物种之间普遍存在的一种相互关系，其中物种利用环境中其他成员的代谢产物以促进自身生长的情形称为代谢互养关系，这种关系对物种间的竞争结果往往有很大影响，甚至会改变种群结构。为了研究代谢互养关系在维持微生物物种多样性中的作用，构建包含不同代谢互养关系的资源竞争模型，这些模型既体现了微生物物种竞争资源时种群密度及资源量的动态，也展示了物种利用其他竞争者的代谢资源对自身生存状况的影响。数值模拟结果显示：（1）考虑微生物中不同的代谢互养关系结构：两物种间单向互养、双向互养以及多物种间的互养，不同的互养关系都可以促进竞争物种稳定共存，竞争中处于劣势的物种通过利用其他竞争成员的代谢产物，打破外界资源量对其生长的限制，改变原本消亡的命运；而处于优势的物种则通过利用其他竞争成员的代谢产物，增大种群密度。（2）多物种竞争同一种有限资源时，不是所有物种都能共存，在四物种模拟中，原本处于最劣势的物种灭绝，其余三者共存。物种产生代谢资源对其本身是"不利"的，如果在模拟中物种利用代谢资源的能力相同，那么物种竞争外界资源的劣势就很可能无法被抵消。通过改变资源利用率发现只有互养关系中代谢资源的利用可以弥补劣势种在竞争外界资源时的不足，多物种才可以全部共存。（3）验证数值模拟结果的普遍性，分析参数变化对共存的影响，结果表明代谢互养关系促进的共存对代谢资源相关参数不敏感，参数的改变只影响平衡态时物种的种群密度。所以，代谢互养关系可以促进相互竞争的微生物物种共存，即微生物之间的互养关系很可能是维持物种多样性的一种机制。     

物种对资源竞争的动力机制及数值模拟

建立了集合种群物种在两个斑块中对资源竞争的数学模型,并进行了数值模拟实验,结果表明：（1）通过R^＊来预测竞争物种的结局,存在几种可能性：一是具有低R^＊值的物种竞争取代高R^＊值的物种;二是具有不同R^＊值的物种,甚至是具有相同R^＊值的物种也存在共存的可能性;三是具有高R^＊值的物种也可以竞争排斥低R^＊值的物种,结论存在不确定性。（2）竞争物种的随机迁移形成了源一汇结构,对物种竞争共存具有促进作用,但弱的资源利用者（较高的R^＊）的迁移率不宜过高。（3）在种群统计率相同的条件下,资源增长率差异越大,越不利于消费者物种的共存;若种群统计率不相同,在资源增长率相同的情况下,物种共存又是不可能的,在自然界中,物种共存需要资源增长率的差异。（4）不同类型的资源增长对竞争物种的稳定性的影响是不同的。     

生物间非传递性竞争研究进展

生物间的竞争关系是决定群落中物种共存和生物多样性的关键因素。传统研究主要关注物种两两之间的竞争作用, 而对多物种相互竞争形成的网络研究相对较少。近年来, 类似于“石头-剪刀-布”游戏的非传递性竞争被认为是一种重要的物种共存和生物多样性的维持机制, 越来越受到生态学家们的关注。本文首先回顾了非传递性竞争定义的发展过程, 并介绍了非传递环的不同结构。其次介绍了基于竞争结局矩阵以及入侵增长率的非传递性竞争度量指标, 并对比不同指标的特点与适用情形。随后通过多个研究实例介绍了非传递性竞争在自然群落中的普遍性, 并指明物种之间的权衡是非传递性竞争产生的生物学机制。最后介绍了非传递性竞争对生物多样性与生态系统功能的影响。非传递性竞争本质上是物种两两之间相互作用的组合, 是只包含单一作用类型的特殊网络结构。因此, 非传递性竞争如何影响生物多样性-生态系统功能关系和群落稳定性, 如何受到环境与高阶相互作用的影响, 以及如何将竞争网络拓展到包含不同相互作用类型的生态网络, 将是未来非传递性竞争研究的重要方向。对非传递性竞争的研究有助于整合生物间的各种相互作用, 构建更加现实合理的生态网络, 并加深对物种共存和生物多样性维持机制的认识, 进而有助于指导生物多样性的保护和恢复工作。     

高阶相互作用对宿主-寄生群落动态的影响

物种间相互作用是影响生物群落稳定性和多样性的重要因素。基于Lotka-Volterra竞争模型,通过构建多宿主种群的种内和种间高阶相互作用模型,研究宿主种群的间接竞争效应对寄生群落动态的影响机制。为有效地揭示高阶作用对种群动态的影响,通过对比宿主-寄生群落的现象模型以及机制模型,利用机制模型产生的合理数据集对现象模型中高阶项的参数进行拟合,进而探讨了高阶相互作用在群落动态中的作用。结果显示,完整的高阶相互作用模型在描述多宿主-寄生系统的群落动态中表现最优,而直接相互作用模型对群落动态的描述相对较差,即同时考虑种间和种内的高阶相互作用模型更加符合机制模型所描述的群落动态。此外,种内高阶作用和种间高阶作用产生不对称效应,宿主间的种间高阶作用对群落产生的影响较种内高阶作用更为显著。该研究结果在一定意义上丰富了宿主-寄生生物群落的稳定性研究,为理解物种间相互作用的多样性研究提供了依据。     

不同栖息地状态下物种竞争模式及模拟研究与应用

物种竞争是影响生态系统演化的重要生态过程之一.而物种在受人类影响出现不同程度毁坏的栖息地上的演化又是非常复杂的,因此研究物种演化对栖息地毁坏的响应是非常必要的.在Tilman研究工作的基础上,将竞争系数引入集合种群动力模式,建立了多物种集合种群竞争共存的数学模型,并对5-物种集合种群在不同栖息地状态下的竞争动态进行了计算机模拟研究.结果表明：（1）不同结构的群落（q值不同）,物种之间的竞争排斥作用强度不同,优势物种明显的群落,物种之间的排斥强度大;（2）随着栖息地毁坏程度的增加,对优势物种的负面影响逐渐减小,而对弱势物种的负面影响逐渐增加;（3）随着栖息地恢复幅度的增加,优势物种和弱势物种之间的竞争越强烈,优势物种受到的竞争排斥加大,而弱势物种逐渐变强,出现了强者变弱、弱者变强的格局;（4）物种竞争排斥与共存受迁移扩散能力和竞争能力影响很大,竞争共存的条件是其竞争能力与扩散能力呈非线性负相关关系;（5）竞争共存的物种的强弱序列发生了变化.     

外来物种入侵后的多物种竞争共存的集合种群模型

基于多物种竞争共存模型,提出外来物种与本地物种竞争共存途径的两种假想:外来物种通过插队竞争与本地物种实现共存;外来物种通过等位竞争与本地物种实现共存.并提出根据外来物种在两种竞争共存模式下占据生境斑块比例的稳定值大小来判断外来物种和本地物种的竞争共存途径.根据两种假想,分别建立了外来物种插队竞争共存模型和等位竞争共存模型.通过应用数学软件Mathematica 4.0对两个模型进行了模拟,得出以下结论:在外来物种与本地物种竞争共存状态下,如果外来物种通过插队竞争与本地物种实现共存,当本地物种竞争力差异较大时,外来物种极易对本地稀少物种构成危害.虽然外来物种不会直接造成本地稀少物种的灭绝,但是会使本地稀少物种的生境斑块急剧减少,增加本地稀少物种灭绝的可能性,而当本地物种竞争力差异较小时,外来物种对本地所有物种的影响都较小.如果外来物种通过等位竞争与本地物种实现共存,无论本地物种竞争力差异大小与否,外来物种只是影响到与其生态位相同的本地物种,影响程度取决于外来物种侵入时所占据生境斑块的比例大小.     

生态位构建对具有资源竞争物种的进化分布动态的影响

生态位构建是有机体通过自身活动、代谢等行为调节或影响其环境或其他生物有机体的过程, 进而导致有机体在自然选择过程中产生不同的进化轨迹。已有研究结论是基于双位点种群遗传模型、基于个体模型以及及元胞自动机模拟模型得出的。目前分析物种进化入侵的常用方法之一是进化分布动态模型。通过引入该方法,着重研究了依特征调节的生态位构建作用对具有种内资源竞争的物种进化动态的影响。结果表明: 依特征调节的生态位构建确实能够引起物种进化轨迹的明显改变。若生态位构建中心位于特征值较小的特征时, 将减少进化分支, 且相较于不存在生态位构建的情况而言, 种群密度提高; 与此相反, 生态位构建中心为特征值较大的特征时,生态位构建维持进化多样性, 尽管其破坏了原有进化的对称分布。这意味着生态位构建通过与环境长期的正反馈作用, 使具有较强依赖的种群特征出现明显差异, 分支数虽有减少, 但显著提升了优势特征的种群密度, 且物种进化分支数随着构建强度的增加而减少。强烈的种内竞争和较弱的生态位构建作用将有利于维持种群的进化多样性。     

物种共存理论研究进展

群落内的多物种如何共存是群落生态学的核心研究内容之一。经典的物种共存理论强调物种之间的生态位分化,注重具体共存机制的研究。这种以具体共存机制为研究对象的方法一定程度上促进了当代物种共存理论框架的形成。在当代物种共存理论框架下,物种间的差异被划分为两类综合性的抽象差异——生态位差异和平均适合度差异,前者促进物种共存,对应稳定化机制;后者导致竞争排除,对应均等化机制。本文在简要回顾经典物种共存理论的基础上,介绍了当代物种共存理论的框架(包括理论的形成和定义)、基于该理论的部分实验验证工作及其在一些重要生态学问题中的应用。当代物种共存理论不仅揭示了群落内物种是如何共存的这一基本理论问题,更重要的是在全球变化的背景下该理论对生物多样性的保护和管理具有重要的应用价值。期望本文的介绍有助于国内生态学和生物多样性工作者了解当代物种共存理论,并将其应用于群落构建和生物多样性维持机制等方面的研究。     

禾-豆混播草地种间竞争与共存

以羊草分别与沙打旺、兴安胡枝子、花苜蓿、紫花苜蓿、山野豌豆5种豆科牧草在混播数量比为1:0、2:1、1:2、0:1的条件下建立两物种混播草地,以相对产量、相对密度和相对产量总值为指标,比较各个混播草地中种间竞争的相对激烈程度;各个物种组合的种间竞争优势以及是否发生氮素资源分离;并探索不同禾-豆混播群落达到共存状态的可能途径.研究结果表明,各个禾-豆组合的相对产量总值分别在不同收获时期大于1,禾草与豆科牧草的生态位发生了不同程度的分离.沙打旺和紫花苜蓿对羊草具有显著的竞争优势,即使其种内竞争大于种间竞争时,混生的羊草亦受到强烈的种间竞争压力.与此相反,羊草对兴安胡枝子、花苜蓿和山野豌豆具有种间竞争优势.刈割对竞争双方的优劣地位产生很大影响,减少强竞争力物种的混播比例,可促进混播物种双方均受益,形成共存格局.实验采用的相对密度指标在预测未来混播种群组成上比相对产量更为可行,并且具有维持低个体大小、高构件密度能力是竞争关系中忍耐型物种能够长期存在的可能原因之一.     

基于结构方程模型分析林分空间结构对草本物种多样性的影响

林分空间结构的改变直接影响林下草本物种的多样性。以针叶纯林、针阔混交林和常绿阔叶林为研究对象,采用结构方程模型研究了林分空间结构对林下草本物种多样性的影响,并探讨了林分水平空间结构,垂直空间结构以及林木竞争态势对林下草本物种多样性的影响的相对重要性。结果表明,林分水平空间结构对林下草本的物种多样性指数和物种均匀性指数均存在极显著的影响（P < 0.001）,影响系数高达0.96和0.89,对林下草本物种丰富度存在显著影响（P < 0.01）,影响系数为0.22;林分垂直空间结对林下草本丰富度和物种均匀性指数均存在极显著的影响（P < 0.001）,影响系数分别为0.86和0.43,对林下草本物种多样性指数存在显著影响（P < 0.01）,影响系数为0.16;林木竞争指数与林下草本丰富度和物种多样性指数也均存在极显著的影响（P < 0.001）,但影响系数较小,分别为-0.47和-0.30,而对林下草本物种均匀性指数未达到显著影响作用（P > 0.05）,影响系数仅为-0.04。整体上看,林分水平空间结构、垂直空间结构和林木竞争态势均对林下草本物种多样性有较强的影响作用,但从影响系数看,林分水平空间结构的影响作用最大,垂直空间结构次之,林木竞争态势的影响作用最小。因此,欲维持或改善林下草本物种多样性,应采取调整林分水平空间结构为主,垂直空间结构调整为辅,并适当降低林木竞争程度的综合经营措施。     

Temporal resource partitioning mitigates interspecific competition and promotes coexistence among insect parasites

A key to understanding life's great diversity is discerning how competing organisms divide limiting resources to coexist in diverse communities. While temporal resource partitioning has long been hypothesized to reduce the negative effects of interspecific competition, empirical evidence suggests that time may not often be an axis along which animal species routinely subdivide resources. Here, we present evidence to the contrary in the world's most biodiverse group of animals: insect parasites (parasitoids). Specifically, we conducted a meta-analysis of 64 studies from 41 publications to determine if temporal resource partitioning via variation in the timing of a key life-history trait, egg deposition (oviposition), mitigates interspecific competition between species pairs sharing the same insect host. When competing species were manipulated to oviposit at (or near) the same time in or on a single host in the laboratory, competition was common, and one species was typically inherently superior (i.e. survived to adulthood a greater proportion of the time). In most cases, however, the inferior competitor could gain a survivorship advantage by ovipositing earlier (or in a smaller number of cases later) into shared hosts. Moreover, this positive (or in a few cases negative) priority advantage gained by the inferior competitor increased as the interval between oviposition times became greater. The results from manipulative experiments were also correlated with patterns of life-history timing and demography in nature: the more inherently competitively inferior a species was in the laboratory, the greater the interval between oviposition times of taxa in co-occurring populations. Additionally, the larger the interval between oviposition times of competing taxa, the more abundant the inferior species was in populations where competitors were known to coexist. Overall, our findings suggest that temporal resource partitioning via variation in oviposition timing may help to facilitate species coexistence and structures diverse insect communities by altering demographic measures of species success. We argue that the lack of evidence for a more prominent role of temporal resource partitioning in promoting species coexistence may reflect taxonomic differences, with a bias towards larger-sized animals. For smaller species like parasitic insects that are specialized to attack one or a group of closely related hosts, have short adult lifespans and discrete generation times, compete directly for limited resources in small, closed arenas and have life histories constrained by host phenology, temporal resource subdivision via variation in life history may play a critical role in allowing species to coexist by alleviating the negative effects of interspecific competition.     

Species coexistence under resource competition with intraspecific and interspecific direct competition in a chemostat

Competition theory has developed separately for direct competition and for exploitative competition. However, the combined effects of the two types of competition on species coexistence remain unclear. To examine how intraspecific and interspecific direct competition contributes to the coexistence of species competing for a single resource, we constructed a chemostat-type resource competition model. With general functions for intraspecific and interspecific direct competition, we derived necessary and sufficient conditions (except for a critical case that rarely occurs in a biological sense) that determine the number of stably coexisting species. From these conditions, we found that the number of coexisting species is determined just by the invasibility of each species into subcommunities with a smaller number of species. In addition, using a combination of rigorous mathematical theory and a simple graphical method, we can demonstrate how the stronger intraspecific direct competition facilitates species invasion, leading to a larger number of coexisting species.     

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.     

Interspecific competition and coexistence of the two chrysomelids,Gastrophysa atrocyanea motschulsky andGalerucella vittaticollis baly (Coleoptera: Chrysomelidae), under limited food resource conditions

Field experiments were conducted in order to investigate the mode of exploitation of food resources and the mechanism of coexistence of mixed larval populations of the two chrysomelids,Gastrophysa atrocyanea andGalerucella vittaticollis, under limited food resource conditions. The larval survival rates seemed high enough to assure coexistence when hatchlings of the two species were released in 1∶1 and 1∶3 ratios on a host plant. However, the survival rate became almost nil for both species when a 3∶1 ratio was employed, suggesting asymmetrical interspecific competition. Wasted food consumption was much higher inG. atrocyanea larvae. The population ofG. atrocyanea seemed to be regulated more by intraspecific competition, while on the other hand, the population ofG. vittaticollis was considered to be more likely affected by the interspecific competition withG. atrocyanea, depending on the initial ratio and density of the two species.     

Effects of spatial aggregation on competition,complementarity and resource use

Abstract The spatial distributions of most species are aggregated to varying degrees. A limited number of studies have examined the effects of spatial aggregation on interspecific and intraspecific interactions, generally finding that spatial aggregation can enhance coexistence between species by reducing the capacity for interspecific competition. Less well studied are the effects of spatial aggregation on complementarity (i.e. differences in resource use strategies) and resource use. Our primary hypothesis was that spatial aggregation reduces the complementarity between species owing to: (i) less interspecific interactions as a result of spatial separation; and (ii) less differences between species as a result of phenotypic plasticity. We further postulate that these negative effects of spatial aggregation on complementarity will reduce resource use by the community. Here we test these hypotheses in a pot experiment in which we applied three levels of spatial aggregation to three sets of two‐species mixtures of herbaceous perennial plant species from native grasslands of south‐eastern Australia. Both root and shoot biomass were significantly affected by spatial aggregation, although the nature of these affects depended upon the species involved, and the relative strengths of interspecific versus intraspecific competition. Complementarity between species in the distribution of their green leaves decreased significantly as spatial aggregation increased for one of the species mixtures, providing some evidence in support of our hypothesis that aggregation reduces complementarity through phenotypic plasticity. Spatial aggregation also altered light interception and use of soil moisture resources, although these effects were dependent on the species involved. We suggest that clear effects of spatial aggregation on complementarity and resource use may be obscured by the idiosyncratic way in which neighbour identity influences plant growth and hence plant size, limiting the ability to generalize, at the community level, any underlying effects of spatial pattern on ecological process.     

Population biology of multispecies helminth infection: competition and coexistence

The role that interspecific interactions play in shaping parasite communities is uncertain. To date, models of competition between helminth species have assumed that interaction occurs through parasite-induced host death. To our knowledge, there has been no theoretical exploration of other forms of competition. We examine models in which competition acts at the point of establishment within the host, and at the time of egg production by the adult worm. The models used are stochastic and we allow hosts to vary in their rate of exposure to infective larvae. We derive the Lotka-Volterra model of competition when exposure is homogenous and thus demonstrate that two helminth species cannot coexist on a single limiting resource. We show that coexistence of species is promoted by heterogeneity in host exposure provided that the rates of exposure to the two species are not perfectly correlated, and, if they are positively correlated, provided that the degree of heterogeneity in host exposure is similar for the two competing helminth species. These results are robust to the mechanism of competition.     

Apparent competition and recovery from infection

We use mathematical models to analyse how the recovery rate from infection influences the fitness of a host in a setting of interspecific competition. We show that sub-optimal immunity against pathogens can be advantageous for the host in the presence of cross-species infection. Weaker immunity allows the parasite to be used as a biological weapon, and this increases the fitness of the host relative to a competitor. A parameter region is observed in which the outcome of competition depends on the initial conditions. We extend this model and consider the dynamics in a spatial setting and find that the outcome depends on the migration rate of the host species. At low migration rates, coexistence of the host species is possible across space. For higher migration rates, the host species characterized by a lower recovery rate can invade the territory of its competitor. Finally, we study these dynamics in an evolutionary setting. Although a lower recovery rate from infection can increase the competitive ability of a species, we find that evolution maximizes the recovery rate and minimizes parasite burden. The models presented are related to the concept of apparent competition, and our results are discussed in relation to both theoretical and empirical studies.     

Host age modulates within-host parasite competition

In many host populations, one of the most striking differences among hosts is their age. While parasite prevalence differences in relation to host age are well known, little is known on how host age impacts ecological and evolutionary dynamics of diseases. Using two clones of the water flea Daphnia magna and two clones of its bacterial parasite Pasteuria ramosa, we examined how host age at exposure influences within-host parasite competition and virulence. We found that multiply-exposed hosts were more susceptible to infection and suffered higher mortality than singly-exposed hosts. Hosts oldest at exposure were least often infected and vice versa. Furthermore, we found that in young multiply-exposed hosts competition was weak, allowing coexistence and transmission of both parasite clones, whereas in older multiply-exposed hosts competitive exclusion was observed. Thus, age-dependent parasite exposure and host demography (age structure) could together play an important role in mediating parasite evolution. At the individual level, our results demonstrate a previously unnoticed interaction of the host''s immune system with host age, suggesting that the specificity of immune function changes as hosts mature. Therefore, evolutionary models of parasite virulence might benefit from incorporating age-dependent epidemiological parameters.     

Mutualism can mediate competition and promote coexistence

Mutualistic interactions are not believed to promote coexistence of competitors because mutualisms produce positive feedbacks on abundances whereas coexistence requires negative feedbacks. Here we show that a mutualism between an anemonefish (Amphiprion) and its sea anemone host mediates the effect of asymmetrical competition for space between the anemonefish and another damselfish (Dascyllus) in a manner that fosters their coexistence. Amphiprion stimulates increases in host area, the shared resource, but social interactions cap the number of anemonefish to two adults per host. Space generated by the mutualism becomes differentially available to Dascyllus because the effectiveness of an anemonefish in excluding its competitor declines with increases in the area it defends. This alters Amphiprion's ratio of per capita intra‐ to interspecific effects and thus facilitates coexistence of the fishes. This mechanism may be prevalent in nature, adding another major pathway by which mutualism can enhance diversity.     

Accounting for interspecific competition and age structure in demographic analyses of density dependence improves predictions of fluctuations in population size

Understanding species coexistence has long been a major goal of ecology. Coexistence theory for two competing species posits that intraspecific density dependence should be stronger than interspecific density dependence. Great tits and blue tits are two bird species that compete for food resources and nesting cavities. On the basis of long‐term monitoring of these two competing species at sites across Europe, combining observational and manipulative approaches, we show that the strength of density regulation is similar for both species, and that individuals have contrasting abilities to compete depending on their age. For great tits, density regulation is driven mainly by intraspecific competition. In contrast, for blue tits, interspecific competition contributes as much as intraspecific competition, consistent with asymmetric competition between the two species. In addition, including age‐specific effects of intra‐ and interspecific competition in density‐dependence models improves predictions of fluctuations in population size by up to three times.