基于土壤食物网的生态系统复杂性-稳定性关系研究进展

复杂性-稳定性关系是生态学核心问题之一。作为模式食物网,土壤食物网在探索生态系统复杂性-稳定性关系中起了极大的作用。总结了以Moore、de Ruiter、Neutel等为代表的理论生态学家以土壤食物网为工具研究生态系统复杂性-稳定性关系的方法、结论及不足之处,并展望了未来的发展方向。Moore、de Ruiter、Neutel等将土壤食物网功能群生物量数据、土壤食物网Lotka-Volterra模型和面向过程模型三者结合起来,描述相互作用强度大小格局、分室、能流组织形式等复杂性特征;将土壤食物网Lotka-Volterra模型与群落矩阵结合起来分析局域稳定性,进而探讨生态系统复杂性-稳定性关系的一般规律。在此基础上,Moore、de Ruiter、Neutel等证明了与随机食物网相比,真实食物网的相互作用强度格局、分室等复杂性特征提高了生态系统稳定性,生产力与稳定性共同决定了食物链的长度,并指出建立在平衡态基础上的静态土壤食物网模型在探索生态系统复杂性-稳定性关系方面具有较大的不足,动态土壤食物网是未来以土壤食物网为工具研究生态系统复杂性-稳定性关系的发展方向。     

土壤微食物网结构与生态功能

土壤微食物网是碎屑食物网中与土壤生态过程密切相关的一部分,通过取食资源基质直接或间接地参与养分循环过程,影响陆地生态系统功能.本文从土壤微食物网的组成、结构和生态功能等方面综述了近年来土壤微食物网的研究进展.通过对土壤微食物网的能量通道及营养级联效应的介绍,阐述了土壤微食物网在碳(C)、氮(N)转化、凋落物分解和植物生长等方面的重要作用.针对目前的研究现状,提出未来土壤生态学研究应与高通量测序及稳定同位素技术相结合;通过构建模型进一步加强对土壤食物网结构和功能的研究,从而深入揭示地下生态过程及其对地上植物生长的反馈作用机理.     

线虫区系分析指示土壤食物网结构和功能研究进展

土壤食物网结构复杂,功能众多,直接测定土壤食物网各功能群生物量并结合数学模型来推断土壤食物网结构和功能,工作量大且分析过程繁琐。线虫生态学的发展为土壤食物网的研究开辟了一条新的思路,即利用线虫区系分析来定性推断食物网的结构和功能。线虫作为土壤中数量最丰富的后生动物,占据着土壤食物网的中心位置,其物种多样性、食性多样性、生活史策略多样性、功能团多样性奠定了其作为土壤食物网结构和功能指示生物的生态学基础。线虫区系分析根据发展历史可以分为个体分类、生活史策略分类、功能团分类和代谢足迹分类四个时期,其中后两个时期主要用于推断土壤食物网结构和功能。基于功能团的线虫区系分析将线虫的食性和生活史策略结合起来,发展出一系列指数来判断土壤食物网的连通性、食物网链长度、外界养分投入情况、分解途径及对外界干扰的响应等。基于代谢足迹的线虫区系分析在功能团分析基础上,加入线虫能流分析,从而定性反映了土壤食物网功能的大小。两者在指示土壤食物网自下而上调节及对植物线虫控制等方面起着重要的作用。     

食物网稳定性机制研究进展:一个基于等级系统的框架

食物网理论沟通了群落生态学和生态系统生态学,将生物多样性和生态系统功能的研究统一起来,是理解生态系统运作机制的关键。自从1973年Robert May的经典研究引发著名的"复杂性-稳定性"论辩之后,人们认识到食物网的稳定性是其结构维持、功能发挥和动态演化的一个重要前提,并开始了对食物网稳定性机制的探索。早期研究主要关注只包含拓扑关系的定性食物网,但后来人们逐渐认识到相互作用强度的重要性,并提出了诸如自限性、弱相互作用、适应性捕食等一系列机制。本文系统梳理了过往研究中模块层面的各类稳定性机制和全网层面对各模块的整合机制,从而清晰地展示了"模块-全网"双层框架的全貌。通过在其基础上的扩展,进而提出了一个基于等级系统的食物网稳定性框架,并从动力学和能量学角度,对各层级内部的稳定性机制以及层级之间的关系进行了探讨,以期为建立普适的食物网稳定性理论提供一些思路。未来的研究方向包括：①将稳定性机制的研究从食物网扩展到更一般的生态网络;②综合考虑生物物理要素、动力学稳定性、系统对能流功率的追求、环境的平稳程度、演化历史等影响因素,从而得到关于食物网结构和动态的更为深刻的认识。     

基于土壤线虫群落分析的油页岩废渣地不同植被类型恢复过程中的土壤食物网能流状况

土壤健康与生态系统功能是当前生态学研究的重要课题。土壤线虫是评价生态系统健康状况的指示生物,特别是用于评价土壤污染和恢复的过程。对广东茂名油页岩废渣场人工实施不同生态恢复改造状况下的土壤线虫属的重要值及土壤线虫反映的土壤食物网能量通道进行了分析,以通过污染地土壤食物网内的能量流动状况反映种植不同植物物种相同时间后的土壤养分与健康状况。结果表明在废渣场种植不同植物物种约8年后,乌墨林和大叶相思林下土壤线虫类群数(属)分别为54和45,重要值最高的是食细菌的棱咽属(Prismatolaimus)和拟丽突属(Acrobeloides);而红荷林和荒草地的线虫类群数分别为41和38,重要值最高的是杂食性的真矛线属(Eudorylaimus);废渣地线虫属的数量最少,仅为34个,重要值最高的是食真菌的丝尾垫刃属(Filenchus)和滑刃属(Aphelenchoides)。食物网能流分析表明:细菌能流通道比重最高的是大叶相思林和乌墨林,而真菌能流通道比重最高的是废渣地,红荷林和荒草地居中,各样地植物能流通道比重都比较小,仅为2%—10%之间。总体来讲,在有植被覆盖的生态系统,养分周转更快,特别是乌墨和大叶相思林,而无植被的废渣地,土壤养分周转速率倾向于更慢的真菌能流通道,这表明在养分条件比较差的情况下,真菌通道的食物网可能会起更大的作用。研究还说明了在油页岩废弃地恢复过程中,植物资源的输入不仅仅刺激植物能流通道,同时也会刺激细菌和真菌能流通道,植被是土壤线虫群落发展的主要驱动力。     

中国两个气候过渡区森林土壤线虫群落的比较研究

为比较不同气候过渡区地带性森林土壤线虫群落结构和多样性,以南亚热带向中亚热带气候过渡区的石门台常绿阔叶林和北亚热带向暖温带气候过渡区的鸡公山落叶阔叶林为样地,探讨不同气候区土壤微食物网能流方式的差异。结果表明,石门台土壤线虫群落属数(S)、生物量(Biomass)、多样性指数(H')、丰富度指数(SR)、成熟度指数(MI)和结构指数(SI)在表层土壤(0~10 cm)均显著高于鸡公山。石门台的土壤线虫通路指数(NCR)均值高于0.5,而鸡公山的NCR均值小于0.5,说明前者土壤食物网可能偏向于细菌能流通道,而后者偏向于真菌能流通道,这也与食细菌线虫、食真菌线虫生物量的计算结果一致。可见线虫群落结构确实存在明显的南北差异,并较好地指示了土壤能流和养分循环状况。     

水生生态系统食物网复杂性与多样性的关系

探索食物网的复杂结构是生态学的中心问题之一。基于构建的黄河口海草床食物网并耦合实际食物网的数据集,整理了包含河口、湖泊、海洋和河流四种水生生态系统类型的48个实际食物网案例。以食物网的节点数反映食物网多样性,物种之间的营养链接数、链接密度和连通度来表示食物网的复杂性,采用营养缩尺模型描述水生生态系统食物网的复杂性特征与节点数的普适性规律。结果表明：所涉及的48个水生生态系统食物网的多样性和复杂性跨度较大,其中,节点数的分布范围为4-124,链接数为3-1830,链接密度为0.75-15.71,连通度为0.06-0.25。不同类型水生生态系统间的连通度存在显著性差异（P=0.01）,节点数、链接数、链接密度不存在显著性差异。各类型生态系统的食物网链接数、链接密度均随节点数的增加而增加（R²=0.92,P<0.001和R²=0.82,P<0.001）。湖泊生态系统的连通度随节点数的变化不明显,围绕在0.20附近;而其他3种类型生态系统的食物网连通度随节点数的增加而降低（R²=0.06-0.41,P<0.001）。对全球尺度的水生食物网多样性和复杂性的定量化研究对于提升对食物网的复杂结构的科学认识,从系统尺度探究多样性和复杂性的关系提供数据支撑。     

群落水平食物网能流季节演替特征

稳定碳、氮同位素比值分析技术是研究生态系统中物质循环与能量流动的有效技术.δ13C可以用来判断食物网中不同生物的能量来源;δ15N主要用于确定生物在食物网中所处的营养位置.通常用δ13C—δ15N图来表征某一特定时间或空间的食物网结构,但是这种方法在比较不同时间和空间食物网结构中功效较差.同时这种定性描述食物网结构也无法满足食物网复杂变化下的假说验证.应用环形统计方法,以太湖梅梁湾鱼类群落为例,定量评价了群落水平食物网能流季节演替特征.结果表明太湖梅梁湾鱼类营养生态位移动的季节特征明显.进一步物种水平分析结果表明,各种鱼类角度和幅度随季节均有变化.Rayleigh检验结果表明,群落中不同种类在秋冬、冬春和夏秋均有显著的一致的方向变化;而春夏期间不显著.Watson-William检验结果表明,群落水平的鱼类营养生态位移动在秋冬和冬春季节转换中有显著差异.引起鱼类群落水平营养生态位在食物网空间中季节性移动的主要因素为可利用资源稳定同位素的季节变化和不同鱼类种群自身的食性季节性转变.同时,由于梅梁湾食物网鱼类群落杂食性水平高,季节性浮游初级生产力成为食物网能量流动的重要驱动作用.因此,在富营养化生态系统中,食物网群落水平营养生态位季节波动也暗示了系统稳定性的下降.定量评价食物网变化有助于认识和理解食物网结构与功能在生态学和生态系统管理等方面的重要.     

农田土壤食物网管理的原理与方法

土壤食物网在维持生态系统生产力和健康等方面起着重要作用,但现代农业中,化肥农药等外部投入已经改变或部分替代了土壤食物网的功能,由此也造成一系列的环境问题。为了协调作物高产与环境保护的利益,需要对土壤食物网进行管理,使土壤食物网符合作物生长的需要,即建立健康土壤食物网。管理土壤食物网有两种方式:(1)直接方式,即通过调节食物网各个功能群的组成来管理土壤食物网;(2)间接方式,即根据农田土壤食物网以自下而上调控方式为主、强调低营养阶层的资源限制的原理,通过调节碎屑的数量和质量来管理食物网。在这两种调控方式中,都需要对被管理的食物网进行监测,监测的方式也分两种,一种是直接测定食物网各功能群的数量和生物量,另外一种方式即以线虫为工具来反应土壤食物网的结构。     

试论土壤线虫多样性在生态系统中的作用

土壤线虫的物种多样性和功能群丰富, 不同功能群的土壤线虫取食习惯也不一样, 而土壤线虫取食习惯的多样性以及它们自身的一些属性决定了它们在土壤中与其他生物之间关系的复杂性。土壤中不同生物之间的相互作用决定了土壤食物网的复杂性, 进而影响着土壤生态系统功能。然而, 有关土壤线虫的多样性与土壤生态系统功能之间的关系的研究还很不足。要全面了解土壤线虫在生态系统中的作用, 研究工作必须: (1) 结合室内模拟和野外控制实验; (2) 结合物种多样性调查和不同功能群的功能分析。     

Trends in research on shallow water food webs

Making sense of the spider–web networks of interactions between species in food webs has been a major pre-occupation of ecologists over the last 60 years. This review describes the early attempts to reduce this complexity through the grouping of individual taxa into functional categories (such as trophic levels), through adopting the energy flow or systems approach as epitomised by the International Biological Programme, and most recently by the derivation of web statistics by food web theorists. The strengths and weaknesses of these approaches are discussed in relation to empirical field experiments for unravelling the processes responsible for organising communities and an assessment made of the representation of these approaches in the marine biological literature.     

Contribution of nematodes to the structure and function of the soil food web

As carbon and energy flow through the soil food web they are depleted by the metabolic and production functions of organisms. To be sustained, a "long" food web, with a large biomass at higher trophic levels, must receive a high rate of rhizodeposition or detrital subsidy, or be top-populated by organisms of slow growth and long life cycle. Disturbed soil food webs tend to be bottom heavy and recalcitrant to restoration due to the slow growth of upper predator populations, physical and chemical constraints of the soil matrix, biological imbalances, and the relatively low mobility and invasion potential of soil organisms. The functional roles of nematodes, determined by their metabolic and behavioral activities, may be categorized as ecosystem services, disservices or effect-neutral. Among the disservices attributable to nematodes are overgrazing, which diminishes services of prey organisms, and plant-damaging herbivory, which reduces carbon fixation and availability to other organisms in the food web. Unfortunately, management to ameliorate potential disservices of certain nematodes results in unintended but long-lasting diminution of the services of others. Beneficial roles of nematodes may be enhanced by environmental stewardship that fosters greater biodiversity and, consequently, complementarity and continuity of their services.     

Intensive agriculture reduces soil biodiversity across Europe

Soil biodiversity plays a key role in regulating the processes that underpin the delivery of ecosystem goods and services in terrestrial ecosystems. Agricultural intensification is known to change the diversity of individual groups of soil biota, but less is known about how intensification affects biodiversity of the soil food web as a whole, and whether or not these effects may be generalized across regions. We examined biodiversity in soil food webs from grasslands, extensive, and intensive rotations in four agricultural regions across Europe: in Sweden, the UK, the Czech Republic and Greece. Effects of land‐use intensity were quantified based on structure and diversity among functional groups in the soil food web, as well as on community‐weighted mean body mass of soil fauna. We also elucidate land‐use intensity effects on diversity of taxonomic units within taxonomic groups of soil fauna. We found that between regions soil food web diversity measures were variable, but that increasing land‐use intensity caused highly consistent responses. In particular, land‐use intensification reduced the complexity in the soil food webs, as well as the community‐weighted mean body mass of soil fauna. In all regions across Europe, species richness of earthworms, Collembolans, and oribatid mites was negatively affected by increased land‐use intensity. The taxonomic distinctness, which is a measure of taxonomic relatedness of species in a community that is independent of species richness, was also reduced by land‐use intensification. We conclude that intensive agriculture reduces soil biodiversity, making soil food webs less diverse and composed of smaller bodied organisms. Land‐use intensification results in fewer functional groups of soil biota with fewer and taxonomically more closely related species. We discuss how these changes in soil biodiversity due to land‐use intensification may threaten the functioning of soil in agricultural production systems.     

Local stability properties of complex,species‐rich soil food webs with functional block structure

Ecologists have long debated the properties that confer stability to complex, species‐rich ecological networks. Species‐level soil food webs are large and structured networks of central importance to ecosystem functioning. Here, we conducted an analysis of the stability properties of an up‐to‐date set of theoretical soil food web models that account both for realistic levels of species richness and the most recent views on the topological structure (who is connected to whom) of these food webs. The stability of the network was best explained by two factors: strong correlations between interaction strengths and the blocked, nonrandom trophic structure of the web. These two factors could stabilize our model food webs even at the high levels of species richness that are typically found in soil, and that would make random systems very unstable. Also, the stability of our soil food webs is well‐approximated by the cascade model. This result suggests that stability could emerge from the hierarchical structure of the functional organization of the web. Our study shows that under the assumption of equilibrium and small perturbations, theoretical soil food webs possess a topological structure that allows them to be complex yet more locally stable than their random counterpart. In particular, results strongly support the general hypothesis that the stability of rich and complex soil food webs is mostly driven by correlations in interaction strength and the organization of the soil food web into functional groups. The implication is that in real‐world food web, any force disrupting the functional structure and distribution pattern of interaction strengths (i.e., energy fluxes) of the soil food webs will destabilize the dynamics of the system, leading to species extinction and major changes in the relative abundances of species.     

重金属污染土壤中生物间相互作用及其协同修复应用

土壤是人类赖以生存的物质基础。我国土壤重金属污染状况不容乐观,给人类健康构成严重威胁。生物修复重金属污染土壤被广泛认为是可持续的修复技术,但当前仍存在修复效率不高的科学瓶颈问题。土壤中生活着丰富的微生物、植物和动物,且这些生物之间存在着复杂的相互作用,并且通过物质循环和能量传递形成了错综复杂的食物网联系。土壤生物间的相互作用能深刻影响土壤中污染物的迁移转化和生物修复的效率,多元生物协同的修复技术集合了单一生物修复方法的优势,具有强化生物修复效果的巨大潜力。文中综述了土壤中微生物-植物-动物之间的相互作用,及其对土壤重金属迁移转化和生物修复效果的影响,并对定向调控土壤食物网结构、提高重金属污染土壤的生物修复效果、建立基于食物网的多元生物协同修复技术进行了展望。     

The relative importance of herbivory and carnivory on the distribution of energy in a stochastic tri-trophic food web

A three-state, discrete-time Markov chain is used to model the dynamics of energy flow in a tri-trophic food web. The distribution of energy in the three trophic levels is related to the rates of flow between the trophic levels and calculated for the entire range of possible flow values. These distributions are then analysed for stability and used to test the idea that plants are resource-limited and herbivores are predation-limited. Low rates of death and decomposition, when coupled with low rates of herbivory and carnivory, tend to destabilize this food web. Food webs with higher rates of death and decomposition are relatively more stable regardless of rates of herbivory and carnivory. Plants are more prone to resource-limitation and herbivores are, in general, limited by their predators, which supports Hairston et al. (Am. Nat. 94 (1960) 421). The rate of decomposition often mediates the roles of top-down and bottom-up control of energy flow in the food web.