生态系统生态学研究的关键问题及趋势——从“整体论与还原论的争论”看

在生态学领域中,存在着生态系统整体论与还原论的争论。Tansley A.G.提出,生态系统是"准有机体"。Odum兄弟提出的"生态系统能量说"被广泛接受,但也受到质疑,称其为"还原论者的整体论"。基于对上述质疑的回应以及对生态系统整体论的追求,Patten B.C.等提出"生态网络理论",运用"网络‘环境子’分析"方法,试图从物理层面分析解决生物层面的"涌现性"问题。不过,这一理论也受到批判,认为其在探究符号化的现象对生态系统的动态影响时,陷入了还原论困境。Jrgensen S.E.等更进一步,提出"系统论"的生态系统生态学,试图从系统科学的角度研究生态系统的"物质-能量-信息-网络"系统。这一理论受到生态学界高度重视,但是也存在着在具体研究过程中如何平衡能量视角和生物地球化学视角的问题。由上述争论可见,生态系统生态学研究的趋势是从"物质实体"到"能量流动",再到"网络信息",最后到"开放系统"层层递进。目前面临的关键问题是:如何在更好地定义生态系统整体性的基础上,采取相应的能够体现生态系统整体性的方法,去获得更多、更好的生态系统整体性的认识。     

Ecosystem growth and development

One of the most important features of biosystems is how they are able to maintain local order (low entropy) within their system boundaries. At the ecosystem scale, this organization can be observed in the thermodynamic parameters that describe it, such that these parameters can be used to track ecosystem growth and development during succession. Thermodynamically, ecosystem growth is the increase of energy throughflow and stored biomass, and ecosystem development is the internal reorganization of these energy mass stores, which affect transfers, transformations, and time lags within the system. Several proposed hypotheses describe thermodynamically the orientation or natural tendency that ecosystems follow during succession, and here, we consider five: minimize specific entropy production, maximize dissipation, maximize exergy storage (includes biomass and information), maximize energy throughflow, and maximize retention time. These thermodynamic orientors were previously all shown to occur to some degree during succession, and here we present a refinement by observing them during different stages of succession. We view ecosystem succession as a series of four growth and development stages: boundary, structural, network, and informational. We demonstrate how each of these ecological thermodynamic orientors behaves during the different growth and development stages, and show that while all apply during some stages only maximizing energy throughflow and maximizing exergy storage are applicable during all four stages. Therefore, we conclude that the movement away from thermodynamic equilibrium, and the subsequent increase in organization during ecosystem growth and development, is a result of system components and configurations that maximize the flux of useful energy and the amount of stored exergy. Empirical data and theoretical models support these conclusions.     

Understand ecosystem regime shifts by modelling ecosystem development using Boolean networks

Complex ecosystems are difficult to model and are still poorly understood. In spite of the long lasting efforts to predict them, we still lack a real integrated framework to grasp their behaviours. In this paper, the concept of ecosystem development is proposed to understand the sharp regime shifts they cumulate over the long term. To handle regime shifts, we develop an integrated model merging the physical, biological and social components and interactions (processes) of an ecosystem into a single graph representation. We then formalize sharp structural (topological) changes of the system with Boolean networks. We illustrate this theoretical approach borrowed from discrete model formalisms on some ecosystems assumed to be representative of most ecosystems: the eusocial (termite and ant) insect colonies and their associated interactions.Boolean networks, combined with rigorous production/rewriting rules and syntax, simulate the ecosystem's responses to strong perturbations. This deterministic and qualitative approach allows identifying fragile components, analysing ecosystem resilience to perturbations and to map out every lethal trajectory of the ecosystem. In our models, termite colonies appear to be almost six times less vulnerable to perturbations than ant colonies, and need (8%) more time steps to collapse. This robust result was partly due to the higher number of essential nodes in ant ecosystems, and partly due to interactions related to the ants' vertebrate predators. The rigorous and parsimonious abilities of such discrete models pave the way in reinterpreting and managing a wide range of ecosystems.     

Flow analysis sensitivities for models of energy or material flow

Sensitivity analyses have been used to examine the flow structure of two hypothetical ecosystem models. These analyses have results which relate to important aspects of ecosystem theory. Cycles are shown to increase the sensitivity of the network, while increased throughflow is shown to decrease the sensitivity. Such results indicate that several factors can be modified to decrease the sensitivity of ecosystems to environmental stress.     

生态系统保护现状及保护等级评估——以江西省为例

基于生态系统的保护是防止生物多样性丧失的重要手段,已成为保护生物学研究的热点。对生态系统保护等级进行划分,确定局域、区域和全球尺度上生态系统保护的优先性,可为制定生态系统保护方案提供重要依据。目前,生态系统保护等级划分的常用指标包括面积流失率、幅度和代表性。以江西省生态系统为例,基于20世纪80年代的江西省植被图、生态系统图、2010年土地利用图和自然保护区图,在GIS环境下进行图层叠加运算和重分类,再进行定量评估和归一化,将江西省生态系统划分为极重要、重要和一般3个级别。结果表明:过去的30年间江西省自然生态系统面积共减少了82613.83km2,减少率超过了60%。全省的37个自然生态系统类型中有19个类型建有国家级自然保护区,但得到保护的面积比例较低。森林生态系统中极重要、重要和一般生态系统类型分别占省国土面积的16.7374%、5.5310%和3.8242%,受国家级自然保护区保护的比例分别为0.51%、3.73%和5.76%;灌丛生态系统中极重要、重要和一般生态系统类型分别占0.0975%、0.9335%和0.0100%,保护比例分别为1.72%、0.17%和0.70%;草地生态系统中极重要、重要和一般生态系统类型分别占0.2647%、0.0005%和0.1064%,保护比例分别为0.21%、0.00%和3.49%;湿地生态系统中极重要、重要和一般生态系统类型分别占0.3532%、0.0345%和1.5650%,保护比例分别为1.87%、0.00%和18.01%。从各级生态系统空间分布来看,极重要生态系统分布范围最广,主要分布在江西省的东南部和西北部;重要生态系统主要分布在西北部和东北部;一般生态系统分布范围最小,主要分布在北部。中部地区的自然生态系统多数由于经济发展而退变为农田、城市等人工生态系统。结合江西省生态系统现状,将极重要的生态系统作为重点保护范围,因而占江西省国土面积17.46%的生态系统应该得到优先保护。这一结果为有效地开展生态系统管理,制定有针对性的生态系统的保护规划具有重要参考价值。     

Assessment of long-term changes of ecosystem indexes in Tongoy Bay (SE Pacific coast): Based on trophic network analysis

Quantitative macroscopic's indexes have been used to compare three trophic models of the exploited benthic ecosystem of Tongoy Bay. In this system the primary productivity and benthic invertebrates are more important in the cycling of biomass. The models were built with a similar number of compartments for the years 1992, 2002 and 2012, using Ecopath with Ecosim (EwE). Odum and Ulanowicz's frameworks and ecological network analysis were then used to estimate the levels of maturity, growth and development of the system. Likewise, “keystoneness” indexes – at each time – were also estimated for the models. Our results show that Tongoy Bay exhibited an increase in maturity and development (“health”) in 2012 compared to past conditions, which was reflected by (1) an increase in the total system biomass, total system throughput, AMI, and absolute Ascendency, (2) higher flow and increased efficiency of transferred energy and its proportion at higher trophic levels, (3) an increase of recycling (FCI), (4) a reduction of NPP/R and NPP/B ratios of the system, and (5) an increase in the number of compartments trophically linked that comprise the keystone species complex. We argue that these results are a consequence of reduced fishing pressure on this benthic system in recent years. This study shows that the fishing would not only have a direct impact on exploited species, but would also affect the structure and functioning of the ecosystem. The information obtained could help to improve the management of fisheries resources, evaluating surveillance indicators that can show the putative changes of intervened ecosystems.     

Trophic networks: How do theories link ecosystem structure and functioning to stability properties? A review

In the context of present global changes, interest in understanding how systems respond to anthropogenic environmental pressures and stress has increased. Indices that characterize ecosystem state are helpful tools for the interpretation of ecosystem responses. The central question is how to link these responses to ecosystem structure and functioning and to quantify ecosystem persistence, resistance or resilience. Quantification and characterization of trophic networks by ecological network analysis (ENA) indices is proceeding rapidly, especially in the field of coastal ecology. In this contribution, we review several theories that relate ecosystem structure and function to stability. The structure and functioning of ecosystems change during the maturation of ecosystems. In the first section, the maturation of ecosystems is described using thermodynamics. In the second and third parts of this paper, we define some concepts for analysing structure and functioning of food webs and discuss their relation to stability. In the last section, we describe three ENA indices and their link to stability. We demonstrate that ENA provides powerful tools for describing local stability, combining quantitative and qualitative concepts. However, it remains incomplete for describing real conservation cases that combine local and global stability.     

生态网络分析方法研究综述

生态网络分析方法是分析生态系统作用关系、辨识系统内在、整体属性的一种有效的系统分析方法。总结了生态网络分析方法的主要研究成果:网络结构特性、网络稳定性、网络上升性、网络效能等;介绍了构建生态网络模型过程和群落构建规则;以德国西部城市诺伊斯河口氮循环为例,介绍David K是如何运用生态网络分析方法来揭示网络中的微动力流循环规律。生态网络分析方法的主要贡献:(1)对人们凭经验感知的生态系统分室间的关联关系,采用了严密的数学模型和推导进行了描述和证明;(2)为生态系统的微动力流循环的研究提供了方法,对生态系统中物质流的间接循环作用进行了科学论证;(3)不仅为分析生态系统提供了一种科学的数学方法,而且,它为探索生态系统提供了不同与牛顿世界观的崭新的认识论。总结与回顾生态网络分析方法,有益于该方法的运用和进一步完善。     

Centrality measures and the importance of generalist species in pollination networks

Studies of complex networks show that nodes with high centrality scores are important to network structure and stability. Following this rationale, centrality measures can be used to (i) identify keystone species in ecological networks, a major issue in community ecology, and (ii) differentiate the keystone species concept, e.g. species may play a key role in a network for different topological reasons. In 34 pollination communities we examine the relationship between the generalization level of species (ND) and two complementary centrality indices: closeness (CC) and betweenness centrality (BC). CC measures the proximity of a species to all other species in the community, while BC describes the importance of a species as a connector. Most networks had a linear ND–CC relationship with a minimum CC value of 0.41. Hence, species were close to each and will be likely to be rapidly affected by disturbances. Contrarily, in most networks, the ND–BC relationships were power-law distributed with exponents larger than one. Only 59% of the species were connectors (BC > 0). In particular, there was a connector threshold value of ND = 0.46. Species above this threshold represent ～40%, almost all of which were connectors. These results indicate that in pollination systems the most generalized species are usually network keystone species, playing at least two roles: (i) interact closely with most other species (high CC) and (ii) connect otherwise unconnected subnetworks (high BC). We discuss the implications of centrality measures to community-based conservation ecology.     

A Bio-Inspired Methodology of Identifying Influential Nodes in Complex Networks

How to identify influential nodes is a key issue in complex networks. The degree centrality is simple, but is incapable to reflect the global characteristics of networks. Betweenness centrality and closeness centrality do not consider the location of nodes in the networks, and semi-local centrality, leaderRank and pageRank approaches can be only applied in unweighted networks. In this paper, a bio-inspired centrality measure model is proposed, which combines the Physarum centrality with the K-shell index obtained by K-shell decomposition analysis, to identify influential nodes in weighted networks. Then, we use the Susceptible-Infected (SI) model to evaluate the performance. Examples and applications are given to demonstrate the adaptivity and efficiency of the proposed method. In addition, the results are compared with existing methods.     

The organisational structure of protein networks: revisiting the centrality–lethality hypothesis

Protein networks, describing physical interactions as well as functional associations between proteins, have been unravelled for many organisms in the recent past. Databases such as the STRING provide excellent resources for the analysis of such networks. In this contribution, we revisit the organisation of protein networks, particularly the centrality–lethality hypothesis, which hypothesises that nodes with higher centrality in a network are more likely to produce lethal phenotypes on removal, compared to nodes with lower centrality. We consider the protein networks of a diverse set of 20 organisms, with essentiality information available in the Database of Essential Genes and assess the relationship between centrality measures and lethality. For each of these organisms, we obtained networks of high-confidence interactions from the STRING database, and computed network parameters such as degree, betweenness centrality, closeness centrality and pairwise disconnectivity indices. We observe that the networks considered here are predominantly disassortative. Further, we observe that essential nodes in a network have a significantly higher average degree and betweenness centrality, compared to the network average. Most previous studies have evaluated the centrality–lethality hypothesis for Saccharomyces cerevisiae and Escherichia coli; we here observe that the centrality–lethality hypothesis hold goods for a large number of organisms, with certain limitations. Betweenness centrality may also be a useful measure to identify essential nodes, but measures like closeness centrality and pairwise disconnectivity are not significantly higher for essential nodes.     

城市生态网络分析研究进展

自生态网络分析方法提出40多年来,其理论发展和应用实践不断拓展,但直至21世纪才不断引入到城市生态系统研究中,用以分析城市内部多个主体和多种生态流构成的关联网络。目前,城市生态网络分析集中于生态网络分析方法与指标的拓展及多尺度的应用研究,而生态网络分析方法又形成了上升性分析和环境元分析两大分支,多尺度应用涵盖了城市镶嵌的区域背景尺度和城市内部产业部门之间的细节尺度。然而,当前研究仍存在着多尺度融合、多种生态网络分析方法集成不足等问题,这限制了城市生态网络分析方法在城市规划设计中的应用。未来城市生态网络分析研究集中于如下3点:(1)开展多尺度城市生态网络分析,包括城市群-城市-园区/社区等,构建多级嵌套生态网络模型;(2)集成上升性与环境元分析方法,提出由外在表征到内在过程的城市生态系统评价模式及模拟方法;(3)强调自然节点在城市生态网络中的重要作用,形成社会经济节点与自然节点并重的生态网络模型,并强调构建多精度的生态网络模型服务于不同的研究目的。     

江西红壤坡地柑橘园生态水文特征及水土保持效益

在江西省水土保持生态科技园对标准柑橘试验小区生态水文特征进行9年的定位观测,设置7个处理,研究柑橘园生态水文特征及水土保持效益.结果表明： 7个处理的平均减流率和减沙率分别为78.5%和77.2%,其中,林下百喜草(Paspalum natatu)带状覆盖、百喜草全园覆盖和水平梯田+梯壁植草减流率较高,分别为94.8%、94.3%和92.5%;柑橘清耕地减流率较低,为33.1%;林下套种黄豆和萝卜减流率居中,为66.0%和77.5%,且横坡耕作优于纵坡耕作.对2009-2010年发生的43场平均降雨量为20.07 mm的雨水柑橘林冠再分配格局进行观测,穿透降〖JP2〗雨量平均为9.15 mm,树干茎流量平均为4.72 mm,林冠截留量平均为620 mm,分别占林外降雨量的44.7%、25.7%和29.6%.随着林外降雨量的增大,林冠层的穿透雨量、茎流量呈递增趋势.当降雨量<10 mm时,树冠截留率与林外降雨之间呈显著的线性负相关;当降雨量>10 mm时,二者之间的相关性不明显.柑橘枯落物的持水率与浸水时间呈对数回归关系,最大持水率达326%.合理的林下植被配置对柑橘果园的水土保持具有重要作用.     

Neighbourhood-scale urban riparian ecosystem classification

Urban riparian ecosystems are now recognized as essential components in determining the vulnerability of nature and human systems to climate change, but still remains uncertainty concerning how to stratify and classify urban riparian landscapes into units of ecological significance at spatial scales appropriate for management. Ecosystem classification presents a tool that allows for quantifying the social and ecological processes of ecosystems so as to shape them into relatively homogeneous management objectives, and provides the potential of offering decision support to urban planners based on urban ecological principles. The purpose of this study is to explore and develop a framework for urban riparian ecosystem classification. The classification framework integrates 9 ecosystem components and 29 spatially-explicit variables that characterized by the biophysical landscape, built environment and human population. Then this framework is applied at the neighbourhood scale in Huangpu River basin, Shanghai city, China, employing hierarchical cluster analysis. The results of the ecosystem classification show that the riparian ecosystems in the Huangpu River could be grouped into three principal categories and these ecosystems are aligned along a gradient of increasing urbanization. We highlight the conservation and payment for riparian ecosystem services delivered in the upstream of Huangpu River, and revegetation and enhancement of riparian ecosystems in the more urbanized downstream area. We also find that riparian vegetation connectivity and coverage were negatively related to land use mix, built environment density, and economic wealth. In terms of riparian vegetation ecology, neighbourhood scale land use mix and built environment density are more influential than socioeconomic background (such as average family income and immigrant status).     

基于信息网络模型的生态风险评价

人类开发活动造成剧烈的生态系统自然条件变化,生态风险评价可以对受到人为干扰下生态系统(包括物种和群落等)的潜在影响进行模拟和量化。通过对信息流量的概念和网络控制分析,综合考虑生态系统组分间的直接和间接作用,提出一种能实现全局风险模拟的生态网络模型,即信息网络模型。在该模型基础上,建立了面向整体生态系统的生态风险评价框架,同时实现兼容多胁迫因子统一模拟和多风险受体间的风险追踪。以澜沧江漫湾水库为例,在估算重金属Hg、Pb和Cd初始环境风险后,利用信息网络模型追踪分析生态系统中不同生态功能组分之间的风险传递路径,评估各生态组分和整体系统的危险程度。结果表明,在累积效应作用下,对于生态系统和部分群落,整合网络风险值与初始环境风险值之间有着显著差别;在发生环境胁迫时,虽然处于食物网底层的生物类群可能最先受险,但在控制信息作用下食物网上层类群也会受险,甚至其最终受到的潜在威胁比前者更大。信息网络模型可识别出复杂的风险流动路径和群落间的风险累积,从而为生态系统风险评价和管理提供更为系统综合的理论依据。     

Functional links and robustness in food webs

The robustness of ecosystems to species losses is a central question in ecology, given the current pace of extinctions and the many species threatened by human impacts, including habitat destruction and climate change. Robustness from the perspective of secondary extinctions has been addressed in the context of food webs to consider the complex network of species interactions that underlie responses to perturbations. In-silico removal experiments have examined the structural properties of food webs that enhance or hamper the robustness of ecosystems to species losses, with a focus on the role of hubs, the most connected species. Here we take a different approach and focus on the role of the connections themselves. We show that trophic links can be divided into functional and redundant based on their contribution to robustness. The analysis of empirical webs shows that hubs are not necessarily the most important species as they may hold many redundant links. Furthermore, the fraction of functional connections is high and constant across systems regardless of size and interconnectedness. The main consequence of this scaling pattern is that ecosystem robustness can be considerably reduced by species extinctions even when these do not result in any secondary extinctions. This introduces the possibility of tipping points in the collapse of ecosystems.     

Evaluating Network Analysis Indicators of Ecosystem Status in the Gulf of Alaska

This is the first study on the emergent properties for empirical ecosystem models that have been validated by time series information. Ecosystem models of the western and central Aleutian Islands and Southeast Alaska were used to examine indices of ecosystem status generated from network analysis and incorporated into Ecopath with Ecosim. Dynamic simulations of the two ecosystems over the past 40 years were employed to examine if these indices reflect the dissimilar changes that occurred in the ecosystems. The results showed that the total systems throughput (TST) and ascendancy (A) followed the climate change signature (Pacific decadal oscillation, PDO) in both ecosystems, whereas the redundancy (R) followed the inverse trend. The different trajectories for important species such as Steller sea lions (Eumetopias jubatus), Atka mackerel (Pleurogrammus monopterygius), pollock (Theragra chalcograma), herring (Clupea pallasii), Pacific cod (Gadus macrocephalus) and halibut (Hippoglossus stenolepis) were noticeable in the Finn cycling index (FCI), entropy (H) and average mutual information (AMI): not showing large change during the time that the Stellers sea lions, herring, Pacific cod, halibut and arrowtooth flounder (Atheresthes stomias) increased in Southeast Alaska, but showing large declines during the decline of Steller sea lions, sharks, Atka mackerel and arrowtooth flounder in the Aleutians. On the whole, there was a change in the emergent properties of the Aleutians around 1976 that was not seen in Southeast Alaska. Conversely, the emergent properties of both systems showed a change around 1988, which indicated that both systems were unstable after 1988.     

The ecological role of cephalopods and their representation in ecosystem models

Cephalopods, especially squids, are believed to have a structuring role in marine ecosystems as a link between different trophic levels, primarily due to their voracious prey consumption and high production rate. Cephalopod ecology, however, is still poorly understood as observational studies often give highly uncertain and variable results due to the peculiarities of cephalopod behaviour and biology, and their responsiveness to external drivers. This review evaluates our representation of cephalopods in ecosystem models and the insights given by these models on the role of cephalopods in our oceans. We examined ecosystem models from 13 regions to analyse the representation of cephalopods and compared their results to local trophic studies. Our analysis indicated that most ecosystem models inadequately include cephalopods in terms of model structure and parametrization; although some models still have the capacity to draw valuable conclusions regarding the impact and role of cephalopods within the system. Oceanic squid species have a major role linking trophic levels and food webs from different habitats. The importance of neritic species varies locally, but generally cephalopods have a substantial impact via their consumer role. To better understand the ecological role of cephalopods, improved representation of these species in ecosystem models is a critical requirement and could be achieved relatively easily to more accurately articulate the mechanisms regulating the ecological role of cephalopods.