浅水湖泊稳态转换预警识别方法局限与展望

浅水湖泊水体底泥交换强烈,极易受人类活动干扰,超过一定阈值即可能发生灾难性的稳态转换,对其有效识别有助于湖泊富营养化的及时防控与修复。浅水湖泊稳态转换可通过系统关键变量(叶绿素、溶解氧、浮游动物、鱼类等)的时间序列(判别不同稳态)、预警信号及阈值等进行识别,其中预警识别可为湖泊生态系统稳态转换提供预判信息,有利于早预警早行动。目前,浅水湖泊稳态转换预警识别因子(方差及自相关性等)主要用于"临界慢化"现象,但在强大外力作用、强烈随机扰动及极端事件下,这些"临界慢化"因子则可能出现误用或错用。基于浅水湖泊基本特征,针对稳态转换的不同驱动机制,探讨"临界慢化"因子的适用性与局限性,并展望其未来发展方向,旨在为湖泊生态系统稳态转换预警识别提供科学参考。     

干旱区生态系统稳态转换及其预警信号——基于景观格局特征的识别方法

在日益加剧的气候变化和土地开垦、放牧等人类活动干扰下,具有多稳态特征的干旱区生态系统可能会经历从相对健康状态到退化状态的稳态转换,导致生态系统的功能下降。早期预警信号的识别是生态系统稳态转换研究的热点,也是管理实践中防止生态系统退化的关键环节。以往预警信号研究聚焦于通用信号如自相关性、方差等统计学指标,然而这些指标对于具有特定机制的干旱区生态系统可能并不适用。基于干旱区景观格局特征所发展起来的空间指标为生态系统稳态转换提供了独特的空间视角,对于理解干旱区生态系统退化过程和机理具有科学意义和实践价值。介绍了干旱区生态系统稳态转换现象及其转换机制;聚焦景观生态学的指标和方法,从空间视角总结基于干旱区景观格局特征的关键预警指标(植被覆盖度、植被斑块形态、植被斑块大小频率分布和水文连通性等),重点剖析这些关键指标的概念、量化方法、识别特征及其实践应用;最后针对指标的优势和局限性对未来的研究方向进行展望,包括发掘潜在景观指标,加强干旱区生态系统变化的多种驱动要素的相互作用机制研究,开展多时空尺度的实证研究,构建生态系统稳态转换预警信号的整体分析框架,以及加强指标阈值的量化研究等方面。     

生态系统的多稳态与突变

许多生态系统可能在短时间内发生难以预料的状态突变, 其中一些生态系统突变的机理可以用多稳态理论进行解释。近年来生态系统的多稳态和突变现象及其机理吸引了研究者和管理者的广泛关注。本文重点对生态系统多稳态的理论基础、识别方法及稳态转换发生的早期预警信号进行综述, 并基于典型生态系统过程对现实世界中可能观测到的稳态转换进行实例分析, 最后对多稳态概念框架和理论应用中的潜在争议进行讨论, 以期为非线性生态系统动态的理论研究、管理实践和生物多样性保护等提供参考。     

草藻型稳态转换对湖泊微生物结构及其碳循环功能的影响

湖泊是地球表层系统中水、土、气等各个圈层相互作用的联结点,对区域物质如碳等元素循环具有重要影响.微生物是湖泊等水生态系统中的重要组成部分,是湖泊等生态系统中碳等元素物质循环的主要驱动者,是深入了解湖泊碳循环过程的关键.受人类活动等影响,湖泊生态系统,尤其是浅水湖泊生态系统往往表现出以高等水生植物(草型)为主要初级生产者的清水稳定态和以浮游藻类(藻型)为主要初级生产者的浊水稳定态,而随着湖泊营养负荷和湖泊环境条件的变化,这两个不同的稳定态之间可以发生转换或者剧变,这种剧变不仅影响湖泊生态系统中的微生物结构,而且对湖泊中有机碳的形成、循环过程及其微生物驱动机制产生重大影响.本文重点就湖泊生态系统中有机碳的转换与微生物关系以及草藻型稳定态的转换对微生物结构及其碳循环功能的影响等进行综述,进一步分析其中的关键科学问题,以期为深入了解湖泊生态系统中碳等元素循环的微生物驱动过程与机制提供帮助.     

稳态转换视角下生态系统服务变化过程与作用机制

优化生态系统服务供给是实现人类社会与自然生态系统和谐发展的必然途径。生态系统在平衡与非平衡之间复杂的转化模式使生态系统服务研究备受阻力,如何科学地解析生态系统服务内在调控机制是实现从自然资源利用到生态系统功能优化的关键。从论述生态系统稳态转换驱动机制入手,阐明扰动发生后稳态转换的路径、生态系统功能对扰动的响应模式;基于稳态转换视角深入诠释生态系统服务内涵及变化过程,以"结构-过程-功能-服务-人类福祉-可持续性"为核心架构来发展生态系统服务理论框架,并从生态系统敏感性和恢复力等内在属性探讨生态系统服务对结构和功能变化的响应情况;解析当土地利用变化超过生态系统阈值时,各项生态系统服务间的互馈作用。基于稳态转换视角评述生态系统服务变化过程与作用机制,以期为生态系统服务研究及生态系统管理提供新视角。     

生态系统稳态转换检测研究进展

在气候变化、人类活动等影响下,生态系统结构和功能可能发生大规模的突变,导致生态系统从一个相对稳定的状态进入另一个稳定状态,这种现象称为稳态转换。由于生态系统的复杂性,准确刻画生态系统多稳态并界定其临界点尚存在挑战,提升对生态系统稳态转换的检测和预测能力依旧是生态学领域研究的热点和难题。基于多稳态理论和稳态转换经典概念框架,阐释了稳态转换检测的理论基础;归纳总结出四种稳态转换检测方法的原理和优劣势;鉴于稳态转换的尺度依赖性,梳理了单一生态系统、区域综合生态系统和全球生态系统不同尺度下的稳态转换检测方法、研究思路和应用案例。基于研究进展和问题现状,提出在未来研究中,亟待发展适应复杂系统的综合检测方法;创新稳态转换多尺度分析的技术方法体系;深化生态系统稳态转换驱动机制研究,构建多元耦合机理模型;进而深化稳态转换检测结果链接生态系统管理的实践研究;解析生态系统服务和可持续发展机制。     

生态阈值:概念、方法与研究展望

生态阈值概念是20世纪70年代提出的,主要指生态系统的几个稳态之间突然改变的点或区域。在阐明生态系统结构与功能的关系、构建区域可持续发展范式以及服务于生态系统管理和生态红线的划定中,生态阈值的检测和量化有着重要的理论和实践意义。该文首先梳理了前人关于生态阈值的概念、类型的一些提法,从预警研究角度提出可以从两个层次理解生态阈值概念:生态阈值点是系统从量变到质变的转折点,类似于红色界限;而生态阈值带可以理解为量变过程中不同稳态之间的转换区域,类似于黄色与橙色预警边界带。黄色生态阈值表示生态系统可通过自身的调节能力重新达到稳定状态;橙色生态阈值表示需要排除干扰因子使得生态系统重新达到平衡;而红色生态阈值为关键阈值点,超过此阈值,生态系统将发生不可逆的退化甚至崩溃。该文还总结了目前确定生态阈值的主要方法,主要是基于野外观测数据的统计分析与模型模拟方法。最后,基于生态系统服务、生物多样性保护与生态系统管理等几个当今生态学热点研究领域,简单总结归纳了生态阈值的研究现状,并提出生态阈值未来的3个研究难点和方向:1)开展针对生态阈值检测和量化的研究;2)关注生态阈值的尺度效应并加强野外观测;3)发挥生态阈值的预警作用,指导"生态红线"的划定和生态系统管理。     

我国生态系统生态阈值研究基础

生态系统中广泛存在非线性变化,表现为系统状态随着压力的逐渐增加而发生骤然转变。为解释这种变化,国外生态学家提出了生态阈值和稳态转换的概念,不断完善理论和方法体系,开展机理和案例研究,深化对复杂系统演化机制的理解,并开始应用于环境管理。在我国,近几十年来在各类生态系统中开展了大量关于压力-响应的定量化研究,取得了丰富成果。这些研究在本质上与生态阈值和稳态转换理论范式紧密关联。本文以“中国生态阈值和稳态转换案例数据库”为基础,按照河流、湖泊、湿地、森林、草地、河口与海洋、农田、荒漠、城市、冻原10种生态系统类型,筛选归纳了相关生态阈值,并阐释了稳态转换机理。将研究案例与生态阈值、稳态转换理论范式进行衔接,目的是整合多领域研究成果,作为生态系统复杂性研究的基础,推动其在生态环境监测、生态安全预警以及生态标准创新发展领域中的应用。     

生态系统灾难性突变研究进展

生态系统灾难性突变是指一个具有多稳态的生态系统, 当驱动因子临近临界点时, 外界环境条件发生微小改变生态系统则产生剧烈响应, 使生态系统从原有稳态转变为另一种生态效益大大降低的稳态。灾难性突变的前提是多稳态的存在, 多稳态是生态系统在相同条件下, 存在结构和功能截然不同的稳定状态。文章阐述了生态系统中灾难性突变的概念, 综述了灾难性突变在湖泊、海洋、森林等生态系统中的存在性和突变的研究现状, 对灾难性突变过程中的多稳态、恢复力、迟滞效应和灾变的预警现状进行了探讨, 并对灾变的产生机制、恢复力的定量研究、生态系统修复的实践和预警信号的深入研究方面进行了展望。大量的理论研究证实灾难性突变是生态系统中的一个普遍现象, 但从实验角度来看研究成果较少且需不断完善。     

生态系统多稳态研究进展

在多稳态的生态系统中，外力可能导致生态系统状态突然之间发生不可逆转的转变，从而达到一个新的平衡状态。但目前对多稳态理论的系统研究很少，如何使用预警信号来预测生态系统的状态转变依旧是个难题。通过多稳态理论的梳理提出了一个更加综合的多稳态定义，并以放牧模型为例，系统总结了多稳态理论的相关概念，将多稳态理论应用在生态系统演替和扰沌理论的解释中；通过对生态系统稳态转换预警信号的原理、优缺点和应用条件的分析，对不同尺度下多稳态的研究方法进行了归纳；最后提出了目前多稳态领域的研究问题和未来的研究重点。结果表明：（1）将时间和空间预警信号结合在一起，并量化正确预警信号的概率，对错误预警信号的比例进行加权，可能会提供更准确的稳态转换的预报。（2）定量观测试验适用于小尺度的研究，而较大尺度的研究则采用简化的模型来模拟研究，选择正确的尺度极有可能改变预警信号的可靠性。（3）结合多稳态理论研究生态系统临界转换和反馈控制机制，并将基于性状的特征指标和进化动力学纳入其中，是生态系统修复实践的重要研究方向。（4）将多稳态相关理论和生态保护管理政策的实践相结合，是多稳态理论未来应用的前景。本研究为多稳态理论和实践的深入研究提供科学支撑。     

Hydrological regulation drives regime shifts: evidence from paleolimnology and ecosystem modeling of a large shallow Chinese lake

Quantitative evidence of sudden shifts in ecological structure and function in large shallow lakes is rare, even though they provide essential benefits to society. Such ‘regime shifts’ can be driven by human activities which degrade ecological stability including water level control (WLC) and nutrient loading. Interactions between WLC and nutrient loading on the long‐term dynamics of shallow lake ecosystems are, however, often overlooked and largely underestimated, which has hampered the effectiveness of lake management. Here, we focus on a large shallow lake (Lake Chaohu) located in one of the most densely populated areas in China, the lower Yangtze River floodplain, which has undergone both WLC and increasing nutrient loading over the last several decades. We applied a novel methodology that combines consistent evidence from both paleolimnological records and ecosystem modeling to overcome the hurdle of data insufficiency and to unravel the drivers and underlying mechanisms in ecosystem dynamics. We identified the occurrence of two regime shifts: one in 1963, characterized by the abrupt disappearance of submerged vegetation, and another around 1980, with strong algal blooms being observed thereafter. Using model scenarios, we further disentangled the roles of WLC and nutrient loading, showing that the 1963 shift was predominantly triggered by WLC, whereas the shift ca. 1980 was attributed to aggravated nutrient loading. Our analysis also shows interactions between these two stressors. Compared to the dynamics driven by nutrient loading alone, WLC reduced the critical P loading and resulted in earlier disappearance of submerged vegetation and emergence of algal blooms by approximately 26 and 10 years, respectively. Overall, our study reveals the significant role of hydrological regulation in driving shallow lake ecosystem dynamics, and it highlights the urgency of using multi‐objective management criteria that includes ecological sustainability perspectives when implementing hydrological regulation for aquatic ecosystems around the globe.     

Rising variance: a leading indicator of ecological transition

Regime shifts are substantial, long-lasting reorganizations of complex systems, such as ecosystems. Large ecosystem changes such as eutrophication, shifts among vegetation types, degradation of coral reefs and regional climate change often come as surprises because we lack leading indicators for regime shifts. Increases in variability of ecosystems have been suggested to foreshadow ecological regime shifts. However, it may be difficult to discern variability due to impending regime shift from that of exogenous drivers that affect the ecosystem. We addressed this problem using a model of lake eutrophication. Lakes are subject to fluctuations in recycling associated with regime shifts, as well as fluctuating nutrient inputs. Despite the complications of noisy inputs, increasing variability of lake-water phosphorus was discernible prior to the shift to eutrophic conditions. Simulations show that rising standard deviation (SD) could signal impending shifts about a decade in advance. The rising SD was detected by studying variability around predictions of a simple time-series model, and did not depend on detailed knowledge of the actual ecosystem dynamics.     

Theoretical basis for predicting climate-induced abrupt shifts in the oceans

Among the responses of marine species and their ecosystems to climate change, abrupt community shifts (ACSs), also called regime shifts, have often been observed. However, despite their effects for ecosystem functioning and both provisioning and regulating services, our understanding of the underlying mechanisms involved remains elusive. This paper proposes a theory showing that some ACSs originate from the interaction between climate-induced environmental changes and the species ecological niche. The theory predicts that a substantial stepwise shift in the thermal regime of a marine ecosystem leads indubitably to an ACS and explains why some species do not change during the phenomenon. It also explicates why the timing of ACSs may differ or why some studies may detect or not detect a shift in the same ecosystem, independently of the statistical method of detection and simply because they focus on different species or taxonomic groups. The present theory offers a way to predict future climate-induced community shifts and their potential associated trophic cascades and amplifications.     

Early detection of ecosystem regime shifts: a multiple method evaluation for management application

Critical transitions between alternative stable states have been shown to occur across an array of complex systems. While our ability to identify abrupt regime shifts in natural ecosystems has improved, detection of potential early-warning signals previous to such shifts is still very limited. Using real monitoring data of a key ecosystem component, we here apply multiple early-warning indicators in order to assess their ability to forewarn a major ecosystem regime shift in the Central Baltic Sea. We show that some indicators and methods can result in clear early-warning signals, while other methods may have limited utility in ecosystem-based management as they show no or weak potential for early-warning. We therefore propose a multiple method approach for early detection of ecosystem regime shifts in monitoring data that may be useful in informing timely management actions in the face of ecosystem change.     

Changing skewness: an early warning signal of regime shifts in ecosystems

Empirical evidence for large-scale abrupt changes in ecosystems such as lakes and vegetation of semi-arid regions is growing. Such changes, called regime shifts, can lead to degradation of ecological services. We study simple ecological models that show a catastrophic transition as a control parameter is varied and propose a novel early warning signal that exploits two ubiquitous features of ecological systems: nonlinearity and large external fluctuations. Either reduced resilience or increased external fluctuations can tip ecosystems to an alternative stable state. It is shown that changes in asymmetry in the distribution of time series data, quantified by changing skewness, is a model-independent and reliable early warning signal for both routes to regime shifts. Furthermore, using model simulations that mimic field measurements and a simple analysis of real data from abrupt climate change in the Sahara, we study the feasibility of skewness calculations using data available from routine monitoring.     

Regime shifts: catastrophic responses of ecosystems to human impacts

Evidence of abrupt changes in ecosystem states, such as sudden eutrophication in lakes, has been increasingly reported in a variety of aquatic and terrestrial systems. Ecosystems may have more than one state with a self-stabilizing mechanism, so that a shift between states does not occur frequently and is not readily reversible. These big changes are termed regime shifts where often one state is preferred over another. Thus, regime shifts are problematic for ecosystem managers, and the need exists for studies that lead to the identification of thresholds of key variables that trigger regime shifts. Regime shifts are currently difficult to predict and in many cases may be caused by the human pursuit of efficiency in land and water productivity in the last few decades. Here I briefly introduce a theoretical approach to predict the shift between a clear-water state and a turbid state in lakes, the best-studied example of regime shifts. This paper also discusses alternative states in other natural systems besides ecosystems to draw more attention to the research currently being performed on regime shifts. Motomi Genkai-Kato is the recipient of the 10th Denzaburo Miyadi Award.     

A new approach for rapid detection of nearby thresholds in ecosystem time series

Massive changes to ecosystems sometimes cross thresholds from which recovery can be difficult, expensive and slow. These thresholds are usually discovered in post hoc analyses long after the event occurred. Anticipating these changes prior to their occurrence could give managers a chance to intervene. Here we present a novel approach for anticipating ecosystem thresholds that combines resilience indicators with Quickest detection of change points. Unlike existing methods, the Quickest detection method is updated every time a data point arrives, and minimizes the time to detect an approaching threshold given the users’ tolerance for false alarms. The procedure accurately detected an impending regime shift in an experimentally manipulated ecosystem. An ecosystem model was used to determine if the method can detect an approaching threshold soon enough to prevent a regime shift. When the monitored variable was directly involved in the interaction that caused the regime shift, detection was quick enough to avert collapse. When the monitored variable was only indirectly linked to the critical transition, detection came too late. The procedure is useful for assessing changes in resilience as ecosystems approach thresholds. However some thresholds cannot be detected in time to prevent regime shifts, and surprises will be inevitable in ecosystem management.     

Leading indicators of trophic cascades

Regime shifts are large, long-lasting changes in ecosystems. They are often hard to predict but may have leading indicators which are detectable in advance. Potential leading indicators include wider swings in dynamics of key ecosystem variables, slower return rates after perturbation and shift of variance towards lower frequencies. We evaluated these indicators using a food web model calibrated to long-term whole-lake experiments. We investigated whether impending regime shifts driven by gradual increase in exploitation of the top predator can create signals that cascade through food webs and be discerned in phytoplankton. Substantial changes in standard deviations, return rates and spectra occurred near the switch point, even two trophic levels removed from the regime shift in fishes. Signals of regime shift can be detected well in advance, if the driver of the regime shift changes much more slowly than the dynamics of key ecosystem variables which can be sampled frequently enough to measure the indicators. However, the regime shift may occur long after the driver has passed the critical point, because of very slow transient dynamics near the critical point. Thus, the ecosystem can be poised for regime shift by the time the signal is discernible. Field tests are needed to evaluate these indicators.