物种多样性与生态系统功能的关系研究进展

物种的空前丧失促使人们越来越多地开始研究物种多样性与生态系统功能的关系,并探讨其潜在的作用机制.本文根据最新研究进展,归纳了微宇宙实验、"生态箱"实验、Cedar Creek草地多样性实验和欧洲草地实验等代表性实验中关于物种多样性与生产力、稳定性、抗入侵性等生态系统功能的焦点问题,介绍了去除实验在多样性与生态系统功能研究中的应用.在此基础上,提出未来研究所面临的挑战,并对研究趋势进行了展望.主要挑战和趋势有:将小尺度上开展的实验研究扩展到较大的时空尺度上;综合考虑生物因素和非生物因素对多样性变化、生态系统功能的交互作用;营养级之间的相互作用、物种共存机制对物种多样性与生态系统功能关系的影响.     

生产力、可靠度与物种多样性:微宇宙实验研究

近年来,生物多样性与生态系统功能的关系成为生态领域内一个重大科学问题。有一些实验研究表明,物种多样性的降低会使生态系统的生产力、稳定性等功能受损,然而对这些实验结果的解释却产生了激烈的争论,因为有两种机制-“生态位互补”和“抽样效应”都可能会产生这种结果。本项研究通过微宇宙实验探讨了物种多样性与生态系统生产力及其可靠度的关系。在10种单细胞藻类中随机抽取物种,构建具有不同物种丰富度的水生群落,并使同一物种丰富度水平的群落之间没有物种交叉,然后检测物种丰富度对群落生产力及其可靠度的作用,群落生产力以藻类干重表示,自实验开始后第4周起,每周测定1次,共测5次。结果显示：物种丰富度对群落生产力有正效应,并且这种正效应随时间推移而增强;许多混合群落的生产力超过了该群落内所有物种的单产,即发生了超产现象,在实验初期某些特定物种对一些混合群落生产力有主要贡献,而在实验后期却没有任何多物种群落的生产力受个别物种存在与否的影响,群落生产力的可靠度与物种丰富度之间不存在显著相关。从以上结果可以得知：物种多样性对群落生产力有着逐渐增强的正效应;物种多样性对生产力的正效应是生态位互补和抽样效应共同作用的结果,但随时间推移,抽取效应逐渐减弱,本顶研究支持了关于生态位互补与抽样效应在多样性正效应中共同起作用的认识,并说明了这两种机制的相对重要性随时间推移而发生改变。     

生物多样性与生态系统功能:进展与争论

生物多样性与生态系统功能的关系已成为当前人类社会面临的一个重大科学问题,生物多样性的空前丧失,促使人们开展了大量研究工作来描述物种多样性-生态系统功能关系,并试图揭示多样性与系统功能关系的内在机制,本文将多样性对生态系统功能作用机制的有关假说分为统计学与生物学两大类：前者是从统计学角度来解释观察到的多样性-系统功能模式,包括抽样效应,统计均衡效应等;而后者是基于多样性的生物学效应给出的,包括生态位互补,种间正相互作用,保险效应等,本文较为详细地介绍了该领域内有代表性的实验工作,包括“生态箱”实验,Cedar Creek草地多样性实验,微宇宙实验,欧洲草地实验,以及在这些实验结果解释上的激烈争论。     

资源互补效应对多样性——生产力关系的影响

 许多有关物种多样性-生态系统功能关系的观察、理论和实验研究都表明, 在局域尺度范围内, 植物种多样性对生态系统生产力存在正效应。 然而, 对于促成这种关系的潜在生态学机制却缺乏足够的了解。 该实验利用9种一年生栽培牧草, 采用各物种单播及混播的方法, 构建不同多样性梯度的实验群落, 对物种多样性与生态系统生产力的关系及资源互补效应对系统生产力的影响进行了研究。 结果表明, 在一年生植物群落内,植物种多样性在一定程度内对系统生产力存在正效应, 物种多样性与生产力呈二次函数关系, 关系式为y = -98.449x2 + 1 039.2 x - 42.407, (R2 = 0.423 1)。 各物种在资源利用、生长速度和竞争能力等功能特征方面存在较大差异, 最高产物种和最低产物种间产量相差5.8倍。 在同一多样性梯度内, 不同物种组合的群落间生产力和互补效应也存在较大差异, 说明物种的成分对生态系统生产力也有重要影响。 同时,在混播群落中程度不同地存在着资源的互补性利用, 说明物种多样性对系统生产力有增强作用, 但相关分析表明, 互补效应和物种多样性间不存在显著相关关系。互补效应的4种计算方法所反映的资源互补程度有所不同, 每种方法各有利弊, 在对系统的多样性效应作用机制进行评价时, 应根据具体情况, 同时采用几种方法, 以利于对资源互补效应做出恰当的估测。     

生产力与生物多样性关系研究进展

生物多样性与生态系统的功能的关系已经成为人类社会面临的一个重大科学问题。本文简要介绍了生产力与生物多样性的概念和研究背景,综述了生产力与物种多样性关系研究的最新进展与争论：1．是物种多样性还是物种特性或物种组成决定生态系统的功能,目前还没有一致的结论;2．生产力与物种多样性关系的尺度效应极其明显,深刻理解和把握生态学尺度和尺度效应有望成为解析生产力与物种多样性关系的突破口：3．不能只保护所谓的关键种,考虑到生态系统的功能很大程度上依赖于受各式各样物种影响的各种不同的过程,尽最大可能保护最大的多样性,才是谨慎而明智的：4．由于自然生态系统极大的复杂性,生产力与物种多样性关系并没有一般的模式。在对已有成果进行综合分析的基础上,对生产力与物种多样性关系研究中亟待解决的区别物种特性与物种多样性问题、尺度问题、实验设计问题进行了探讨,并对未来的发展方向进行了展望。     

消费者多样性对食物网结构和生态系统功能的影响

前所未有的生物多样性丧失使人们越来越关注生物多样性的生态系统功能.现有的绝大多数研究都是局限在单一营养级别上,主要是植物上,但是今天越来越多的证明表明消费者的多样性对生态系统结构和功能具有深刻影响.综述了消费者多样性对相邻或非相邻营养级的种群密度、物种多样性和生产力等方面影响的最新进展,同时也提出了若干研究展望.总体上.消费者多样性,无论是草食动物还是肉食动物,都倾向于增加该消费者所在营养级的养分和能量利用效率,以及生产力.这可能源于取样效应,或者物种之间的互补作用,类似于植物物种多样性影响初级生产力的机制.草食动物可能降低或者提高植物物种多样性,或者没有显著影响,其具体效应取决于生态系统生产力水平和草食动物的大小.捕食者哌能通过直接抑制草食动物而间接提高植物的多样性和生产力,但这种效应的大小差异很大,甚至效应的方向,都可能随团体内捕食者所占的比例而改变.未来的研究,应该考虑应用较大尺度的实验来检测食物网复杂营养关系对生态系统特性的影响,继续探讨消费者对生态系统功能的影响机制.认为异速生长法则和生态化学计量学在食物网组分关系研究中的应用将有利于增强人们对消费者.生态系统功能关系的理解.另外,全球变暖和转基因植物对食物网中消费者结构和生态系统的功能的影响也将是未来的一个重要研究方向.     

植物遗传多样性与生态系统功能关系的研究进展

全球变化和人类活动导致物种生境的萎缩, 造成很多植物种群数量缩减, 遗传多样性快速丧失。对于物种多样性低的生态系统, 优势种的遗传多样性可能比物种多样性对生态系统功能产生更大的影响。因此, 了解遗传多样性和生态系统功能的关系(GD-EF)及其机制对生物多样性保护、应对环境变化和生态修复具有指导意义。该文综述了植物遗传多样性对生态系统结构(高营养级生物群落结构)和生态系统功能(初级生产力、养分循环和稳定性)的影响及机制、功能多样性对GD-EF的影响、遗传多样性效应和物种多样性效应的比较, 以及GD-EF在生态修复等实际应用的研究进展。最后指出当前研究的不足之处, 以期为后续研究提供参考: 1)还需深入研究GD-EF机制; 2)未评估遗传多样性对生态系统多功能性的影响; 3)不同遗传多样性测度对生态系统功能的影响不明确; 4)缺少长期的和多空间尺度结合的GD-EF实验; 5)遗传多样性效应相对于其他因子的作用不清楚。     

A review on the relationships between plant genetic diversity and ecosystem functioning

全球变化和人类活动导致物种生境的萎缩, 造成很多植物种群数量缩减, 遗传多样性快速丧失。对于物种多样性低的生态系统, 优势种的遗传多样性可能比物种多样性对生态系统功能产生更大的影响。因此, 了解遗传多样性和生态系统功能的关系(GD-EF)及其机制对生物多样性保护、应对环境变化和生态修复具有指导意义。该文综述了植物遗传多样性对生态系统结构(高营养级生物群落结构)和生态系统功能(初级生产力、养分循环和稳定性)的影响及机制、功能多样性对GD-EF的影响、遗传多样性效应和物种多样性效应的比较, 以及GD-EF在生态修复等实际应用的研究进展。最后指出当前研究的不足之处, 以期为后续研究提供参考: 1)还需深入研究GD-EF机制; 2)未评估遗传多样性对生态系统多功能性的影响; 3)不同遗传多样性测度对生态系统功能的影响不明确; 4)缺少长期的和多空间尺度结合的GD-EF实验; 5)遗传多样性效应相对于其他因子的作用不清楚。     

功能多样性对典型阔叶红松林生产力的影响

为比较生物量比率假说与生态位互补假说在解释生产力变异的相对重要性,探讨生物多样性和生产力之间的关系是否受到生物和非生物因素的影响,该研究依托小兴安岭9 hm~2阔叶红松(Pinus koraiensis)林动态监测样地,通过计算群落初始生物量、物种多样性、功能多样性、植物性状的群落加权平均值和测定环境因子,运用线性回归模型、结构方程模型,比较了物种多样性和功能多样性与生产力的相关性。结果表明:(1)物种多样性和功能多样性均对生产力有显著作用,功能多样性比物种多样性与生产力的关系更为密切;(2)功能多样性指数比群落加权平均值能更好地解释生产力变异,说明生态位互补假说更适用于解释阔叶红松林群落内生产力的变异;(3)生物多样性与生产力的关系受生物因素与非生物因素的共同作用,相较于多样性和功能性状组成(植被质量),初始林分生物量(植被数量)能更有效地解释生产力的变异。生物多样性与生产力关系的研究应从植被质量与植被数量同时出发,评估生态系统过程的多种非生物和生物驱动因素,同时维护森林功能多样性,加强植物与土壤环境的保护,对有效增加生产力和维持生物多样性具有重要意义。     

生态系统多功能性的指标选择与驱动因子: 研究现状与展望

全球变化和人类活动正以空前的速度在世界范围内改变着生物多样性, 这导致了全球生物多样性的锐减以及生产力的下降、病虫害的增加和抗入侵能力的减弱等生态问题。近30年来, 生态学家开始对于生物多样性的持续丧失是否以及如何影响生态系统功能的问题越来越感兴趣, 生物多样性与生态系统功能(biodiversity and ecosystem functioning, BEF)关系的研究应运而生, 并成为生态学研究的热点之一。但长期以来, 研究者更多地关注单一生态系统功能, 而忽略了生态系统能够同时提供多种生态系统功能的能力, 即生态系统多功能性(ecosystem multifunctionality, EMF)。本文综述了EMF研究中功能指标的选择、生物多样性的不同维度、微生物多样性对EMF的影响以及其他非生物因子对EMF的驱动等进展。因只考虑单一功能可能会低估生物多样性对整体生态系统功能的影响, 故生物多样性与生态系统多功能性(BEMF)关系的研究成为BEF关系研究的重点。近年来, BEMF关系的研究发展较快, 在不同生态系统(包括水生、草地、森林、旱地、农业等)、不同研究尺度(从区域到全球尺度)、BEMF关系的驱动机制(从单一驱动机制到多种驱动机制共同作用)、研究方法(包括新概念以及新的量化方法的提出和应用)等方面均取得了新的进展。但仍有不足之处, 如对于EMF研究中功能指标的选取没有统一的标准、对地下微生物多样性的关注度不够、涉及多营养级水平下的BEMF关系研究较少、驱动EMF的机制仍存在争论等。未来应加强对于功能指标选取的标准研究, 综合分析地上、地下生物多样性以及非生物因子对EMF的整体影响, 加强生态系统多服务性(ecosystem multiserviceability, EMS)方法的研究和应用。     

Predator diversity and the functioning of ecosystems: the role of intraguild predation in dampening trophic cascades

Single trophic‐level studies of the relationship between biodiversity and ecosystem functioning highlight the importance of mechanisms such as resource partitioning, facilitation, and sampling effect. In a multi‐trophic context, trophic interactions such as intraguild predation may also be an important mediator of this relationship. Using a salt‐marsh food web, we investigated the interactive effects of predator species richness (one to three species) and trophic composition (strict predators, intraguild predators, or a mixture of the two) on ecosystem functions such as prey suppression and primary production via trophic cascades. We found that the trophic composition of the predator assemblage determined the impact of increasing predator species richness on the occurrence of trophic cascades. In addition, increasing the proportion of intraguild predator species present diminished herbivore suppression and reduced primary productivity. Therefore, trophic composition of the predator assemblage can play an important role in determining the nature of the relationship between predator diversity and ecosystem function.     

Density compensation can cause no effect of biodiversity on ecosystem functioning

The role of density compensation (the decline of species density with increasing diversity), in the context of biodiversity and ecosystem functioning, has not been explicitly explored. I used aquatic microbial communities containing bacterivorous consumers (protozoans and rotifers) to investigate whether competition can lead to density compensation and whether density compensation can contribute to the relationship between biodiversity and ecosystem functioning. The experiment employed a nested design in which the consumer diversity gradient (0, 1, 2 or 4 species) was constructed by drawing all possible species or species combinations at each diversity level from a five-species pool. All consumer species coexisted but there was little evidence for overyielding or species dominance, suggesting weak complementarity and sampling effects. Rather, increasing number of consumer species resulted in community-wide density compensation, such that aggregate consumer biomass was unaffected by consumer diversity. Whereas culturable bacterial density declined as consumer diversity increased, total bacterial density showed no discernible response to changes in consumer diversity, a result probably due in part to heterogeneity in bacterial edibility. This study demonstrates the potential for density compensation to shape the relationship between biodiversity and ecosystem functioning.     

A synthesis of subdisciplines: predator–prey interactions, and biodiversity and ecosystem functioning

The last 15 years has seen parallel surges of interest in two research areas that have rarely intersected: biodiversity and ecosystem functioning (BEF), and multispecies predator–prey interactions (PPI). Research addressing role of biodiversity in ecosystem functioning has focused primarily on single trophic‐level systems, emphasizing additive effects of diversity that manifest through resource partitioning and the sampling effect. Conversely, research addressing predator–prey interactions has focused on two trophic‐level systems, emphasizing indirect and non‐additive interactions among species. Here, we use a suite of consumer‐resource models to organize and synthesize the ways in which consumer species diversity affects the densities of both resources and consumer species. Specifically, we consider sampling effects, resource partitioning, indirect effects caused by intraguild interactions and non‐additive effects. We show that the relationship between consumer diversity and the density of resources and consumer species are broadly similar for systems with one vs. two trophic levels, and that indirect and non‐additive interactions generally do little more than modify the impacts of diversity established by the sampling effect and resource partitioning. The broad similarities between systems with one vs. two trophic levels argue for greater communication between researchers studying BEF, and researchers studying multispecies PPI.     

Tree species richness promotes productivity in temperate forests through strong complementarity between species

Understanding the link between biodiversity and ecosystem functioning (BEF) is pivotal in the context of global biodiversity loss. Yet, long-term effects have been explored only weakly, especially for forests, and no clear evidence has been found regarding the underlying mechanisms. We explore the long-term relationship between diversity and productivity using a forest succession model. Extensive simulations show that tree species richness promotes productivity in European temperate forests across a large climatic gradient, mostly through strong complementarity between species. We show that this biodiversity effect emerges because increasing species richness promotes higher diversity in shade tolerance and growth ability, which results in forests responding faster to small-scale mortality events. Our study generalises results from short-term experiments in grasslands to forest ecosystems and demonstrates that competition for light alone induces a positive effect of biodiversity on productivity, thus providing a new angle for explaining BEF relationships.     

Biodiversity and ecosystem functioning: recent theoretical advances

The relationship between biodiversity and ecosystem functioning has emerged as a major scientific issue today. As experiments progress, there is a growing need for adequate theories and models to provide robust interpretations and generalisations of experimental results, and to formulate new hypotheses. This paper provides an overview of recent theoretical advances that have been made on the two major questions in this area: (1) How does biodiversity affect the magnitude of ecosystem processes (short‐term effects of biodiversity)? (2) How does biodiversity contribute to the stability and maintenance of ecosystem processes in the face of perturbations (long‐term effects of biodiversity)?
Positive short‐term effects of species diversity on ecosystem processes, such as primary productivity and nutrient retention, have been explained by two major types of mechanisms: (1) functional niche complementarity (the complementarity effect), and (2) selection of extreme trait values (the selection effect). In both cases, biodiversity provides a range of phenotypic trait variation. In the complementarity effect, trait variation then forms the basis for a permanent association of species that enhances collective performance. In the selection effect, trait variation comes into play only as an initial condition, and a selective process then promotes dominance by species with extreme trait values. Major differences between within‐site effects of biodiversity and across‐site productivity–diversity patterns have also been clarified. The local effects of diversity on ecosystem processes are expected to be masked by the effects of varying environmental parameters in across‐site comparisons.
A major reappraisal of the paradigm that has dominated during the last decades seems necessary if we are to account for long‐term effects of biodiversity on ecosystem functioning. The classical deterministic, equilibrium approaches to stability do not explain the reduced temporal variability of aggregate ecosystem properties that has been observed in more diverse systems. On the other hand, stochastic, nonequilibrium approaches do show two types of biodiversity effects on ecosystem productivity in a fluctuating environment: (1) a buffering effect, i.e., a reduction in the temporal variance; and (2) a performance‐enhancing effect, i.e., an increase in the temporal mean. The basic mechanisms involved in these long‐term insurance effects are very similar to those that operate in short‐term biodiversity effects: temporal niche complementarity, and selection of extreme trait values. The ability of species diversity to provide an insurance against environmental fluctuations and a reservoir of variation allowing adaptation to changing conditions may be critical in a long‐term perspective.
These recent theoretical developments in the area of biodiversity and ecosystem functioning suggest that linking community and ecosystem ecology is a fruitful avenue, which paves the way for a new ecological synthesis.     

Dispersal scales up the biodiversity–productivity relationship in an experimental source-sink metacommunity

The influence of biodiversity on ecosystem functioning is a major concern of ecological research. However, the biodiversity–ecosystem functioning relationship has very often been studied independently from the mechanisms allowing coexistence. By considering the effects of dispersal and niche partitioning on diversity, the metacommunity perspective predicts a spatial scale-dependence of the shape of the relationship. Here, we present experimental evidence of such scale-dependent patterns. After approximately 500 generations of diversification in a spatially heterogeneous environment, we measured functional diversity (FD) and productivity at both local and regional scales in experimental source-sink metacommunities of the bacterium Pseudomonas fluorescens SBW25. At the regional scale, environmental heterogeneity yielded high levels of FD and we observed a positive correlation between diversity and productivity. At the local scale, intermediate dispersal increased local FD through a mass effect but there was no correlation between diversity and productivity. These experimental results underline the importance of considering the mechanisms maintaining biodiversity and the appropriate spatial scales in understanding its relationship with ecosystem functioning.     

Trophic complementarity drives the biodiversity–ecosystem functioning relationship in food webs

The biodiversity–ecosystem functioning (BEF) relationship is central in community ecology. Its drivers in competitive systems (sampling effect and functional complementarity) are intuitive and elegant, but we lack an integrative understanding of these drivers in complex ecosystems. Because networks encompass two key components of the BEF relationship (species richness and biomass flow), they provide a key to identify these drivers, assuming that we have a meaningful measure of functional complementarity. In a network, diversity can be defined by species richness, the number of trophic levels, but perhaps more importantly, the diversity of interactions. In this paper, we define the concept of trophic complementarity (TC), which emerges through exploitative and apparent competition processes, and study its contribution to ecosystem functioning. Using a model of trophic community dynamics, we show that TC predicts various measures of ecosystem functioning, and generate a range of testable predictions. We find that, in addition to the number of species, the structure of their interactions needs to be accounted for to predict ecosystem productivity.     

Trophic and non‐trophic interactions influence the mechanisms underlying biodiversity–ecosystem functioning relationships under different abiotic conditions

Plant diversity effects on ecosystem functioning usually have been studied from a plant perspective. However, the mechanisms underlying biodiversity–ecosystem functioning relationships may also depend on positive or negative interactions between plants and other biotic and abiotic factors, which remain poorly understood. Here we assessed whether plant–herbivore and/or plant–detritivore interactions modify the biodiversity–ecosystem functioning relationship and the mechanisms underlying biodiversity effects, including complementarity and selection effects, biomass allocation, vertical distribution of roots, and plant survival using a microcosm experiment. We also evaluated to what extent trophic and non‐trophic interactions are affected by abiotic conditions by studying drought effects. Our results show that biotic and abiotic conditions influence the shape of the biodiversity–ecosystem function relationship, varying from hump‐shaped to linear. For instance, total biomass increased linearly with plant richness in the presence of detritivores, but not in the absence of detritivores. Moreover, detritivore effects on belowground plant productivity were highly context dependent, varying in the presence of herbivores. Plant interactions with soil biota, especially with herbivores, influenced the mechanisms underlying diversity effects. Herbivores increased plant complementarity and modified biomass allocation and vertical distribution of roots. Furthermore, biotic–abiotic interactions influenced plant productivity differently across plant functional groups. Our findings emphasize the importance of complex biotic interactions underlying biodiversity effects, and that these biotic interactions may change with abiotic conditions. Despite minor changes in productivity in the short‐term, soil biota‐induced changes in plant–plant interactions and plant survival are likely to have significant long‐term consequences for ecosystem functioning. Considering the context‐dependency of multichannel interactions may contribute to reconciling differences among observed patterns in biodiversity studies. Further, abiotic conditions modified the effects of biotic interactions, suggesting that changes in environmental conditions may not only affect ecosystems directly, but also change the biotic composition of and dynamics within ecosystems.     

Interactive prey and predator diversity effects drive consumption rates

The positive relationship between biodiversity and ecosystem functioning is mainly derived from studies concerning primary producers, whereas a generalization of this relationship for higher trophic levels is more difficult. Furthermore, most evidence of the biodiversity–ecosystem functioning relationship is derived from experiments manipulating only one trophic level and, as a consequence, interactive diversity effects at multiple trophic levels have mostly been ignored. Here, we performed a mesocosm experiment in which we manipulated functional group diversity at two trophic levels (primary and secondary consumers) applying a full‐factorial design. More specifically, we asked whether 1) predator functional diversity affects prey mortality rates, 2) prey functional diversity affects prey mortality rates, 3) whether there are interactive effects of simultaneous diversity changes at both trophic levels. For each trophic level we used two functional groups, i.e. organisms belonging to two different habitat domains: at the higher trophic position 1) a ground foraging spider species and 2) a spider species foraging in the vegetation canopy and at the lower trophic position 3) a ground living cricket species and 4) leafhoppers living in the vegetation canopy. Increasing predator functional group diversity increased prey mortality by 53%, and increasing prey functional group diversity increased prey mortality by 24%. Further, prey mortality was highest at the uppermost level of functional group diversity (142% increase in prey mortality compared to single prey and predator functional diversity), most likely due to resource partitioning between the predators. This finding demonstrates that a multi‐trophic perspective is necessary, and that previous studies focusing on only one trophic level have most likely underestimated the strength of the relationship between biodiversity and ecosystem functioning.