A Generic Shallow Lake Ecosystem Model Based on Collective Expert Knowledge

We used fuzzy cognitive mapping (FCM) to develop a generic shallow lake ecosystem model by augmenting the individual cognitive maps drawn by 8 scientists working in the area of shallow lake ecology. We calculated graph theoretical indices of the individual cognitive maps and the collective cognitive map produced by augmentation. There were a total of 32 variables with 113 connections in the collective cognitive map. The graph theoretical indices revealed internal cycles showing non-linear dynamics in the shallow lake ecosystem. The ecological processes were organized democratically without a top-down hierarchical structure. The most central variable in the collective map was submerged plants. The strongest connections were suspended solids concentration decreasing water clarity, phosphorus concentration increasing the phytoplankton biomass, higher water clarity increasing submerged plants, benthivorous fish biomass reducing submerged plants and increasing suspended solids concentration, and submerged plants decreasing suspended solids. The steady state condition of the generic model was a characteristic turbid shallow lake ecosystem. The generic shallow lake ecosystem model had the tendency to go into a turbid state since there were no dynamic environmental changes that could cause shifts between a turbid and a clearwater state, and the generic model indicated that only a dynamic disturbance regime could maintain the clearwater state. The model developed herein captured the empirical behavior of shallow lakes, and contained the basic model of the Alternative Stable States Theory. In addition, our model expanded the basic model by quantifying the relative effects of connections and by extending it with 22 more variables and 99 more weighted causal connections. Using our expanded model we ran 4 simulations: harvesting submerged plants, nutrient reduction, fish removal without nutrient reduction, and biomanipulation. Only biomanipulation, which included fish removal and nutrient reduction, had the potential to shift the turbid state into clearwater state. The structure and relationships in the generic model as well as the outcomes of the management simulations were supported by actual field studies in shallow lake ecosystems. Thus, fuzzy cognitive mapping methodology enabled us to understand the complex structure of shallow lake ecosystems as a whole and obtain a valid generic model based on tacit knowledge of experts in the field.     

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.     

How Regime Shifts in Connected Aquatic Ecosystems Are Affected by the Typical Downstream Increase of Water Flow

All over the world freshwater ecosystems like ponds, ditches and lakes suffer from nutrient-driven regime shifts from submerged plants to dominance by algae or free-floating plants. Although freshwaters are often connected and part of a network, most of our current knowledge on regime shifts comes from studies of isolated ecosystems. The few studies that have assessed the spatial manifestation of regime shifts overlooked the hydrological fact that the water flow through connected waters typically increases in the downstream direction. Here, we use a complex ecosystem model to show that this increase in flow does not lead to spatial differences in ecosystem state. We support these findings with a simple, analytically tractable, nutrient retention model on connected waterbodies. The model shows that all bodies have the same nutrient concentration despite spatial gradients in the flow of water as well as nutrients carried by the water. As a consequence, each connected waterbody is equally vulnerable to a regime shift, implying a regime shift to be system-wide. Furthermore, it appeared that each connected waterbody behaves the same as an isolated waterbody, implying that the vast body of theory on isolated systems, like alternative stable states theory, can still be useful for connected systems. Although these findings are violated when there is heterogeneity in lateral runoff or waterbody characteristics—leading to spatial differences in ecosystem state and therefore to differences in the vulnerability to a regime shift—they show that the typical downstream build-up of water flow does not necessarily lead to differences in ecological state, and thereby provide a basic concept to better understand the ecology of connected freshwaters.     

Contrasting impacts of invasive engineers on freshwater ecosystems: an experiment and meta-analysis

Invasion by common carp (Cyprinus carpio) and red swamp crayfish (Procambarus clarkii) in shallow lakes have been followed by stable-state changes from a macrophyte-dominated clear water state to a phytoplankton-dominated turbid water state. Both invasive carp and crayfish are, therefore, possible drivers for catastrophic regime shifts. Despite these two species having been introduced into ecosystems world-wide, their relative significance on regime shifts remains largely unexplored. We compared the ecological impacts of carp and crayfish on submerged macrophytes, water quality, phytoplankton, nutrient dynamics, zooplankton and benthic macroinvertebrates by combining an enclosure experiment and a meta-analysis. The experiment was designed to examine how water quality and biological variables responded to increasing carp or crayfish biomass. We found that even at a low biomass, carp had large and positive impacts on suspended solids, phytoplankton and nutrients and negative impacts on benthic macroinvertebrates. In contrast, crayfish had a strong negative impact on submerged macrophytes. The impacts of crayfish on macrophytes were significantly greater than those of carp. The meta-analysis showed that both carp and crayfish have significant effects on submerged macrophytes, phytoplankton, nutrient dynamics and benthic macroinvertebrates, while zooplankton are affected by carp but not crayfish. It also indicated that crayfish have significantly greater impacts on macrophytes relative to carp. Overall, the meta-analysis largely supported the results of the experiment. Taken as a whole, our results show that both carp and crayfish have profound effects on community composition and ecosystem processes through combined consequences of bioturbation, excretion, consumption and non-consumptive destruction. However, key variables (e.g. macrophytes) relating to stable-state changes responded differently to increasing carp or crayfish biomass, indicating that they have differential ecosystem impacts.     

Effects of plant harvesting and nutrient enrichment on phytoplankton community structure in a shallow urban lake

The utility of shallow water bodies in urban environments is frequently compromised either by dense beds of submerged plants or cyanobacterial blooms associated with nutrient enrichment. Although submerged plants are often harvested to facilitate recreational uses, this activity may alter the phytoplankton community, which in turn, also may restrict the use of the lake. We tested whether (i) plant harvesting reduced the abundance of flagellate algae and increased the abundance of cyanobacteria, and (ii) whether increasing levels of nutrient enrichment caused shifts in the dominance of heterocytous cyanobacteria, non-heterocytous cyanobacteria and Chlorophyta, in a shallow urban lake in Southern Australia as has been observed for shallow Danish lakes in previous studies. These predictions were tested with large (3000 l), replicated mesocosms in a warm, highly productive, shallow lake densely colonised by the submerged angiosperm, Vallisnaria americana Michaux. The heterokont algae, Chlorophyta, Cyanobacteria and Cryptophyta were the most numerous algal divisions in the lake. The Euglenophyta, although uncommon in early summer, became more abundant towards the end of summer. The Dinophyta and Charophyta were rare. The abundance of the heterokont algae and Euglenophyta was significantly reduced by plant harvesting even after plants had partially re-established 18 weeks after initial harvesting. The decline in the Euglenophyta in response to plant harvesting is consistent with earlier findings, that the relative abundance of flagellate algae tends to be greater in the presence of submerged plants. Contrary to our prediction, we found that the Cyanobacteria did not increase in response to plant harvesting, however the response may be altered under higher nutrient levels. Algal responses to nutrient enrichment in the presence of dense V. americana plants generally followed the patterns observed in shallow Danish lakes despite the large differences in climatic conditions. Both studies found that the abundance of heterocytous cyanobacteria declined at higher levels of nutrient enrichment, whereas non-heterocytous cyanobacteria and chlorophytes increased.     

Palaeoecological evidence for sustained change in a shallow Murray River (Australia) floodplain lake: regime shift or press response?

Paleolimnological techniques can reveal long-term perturbations and associated stable state transitions of lake ecosystems. However, such transitions are difficult to predict since changes to lake ecosystems can be abrupt or gradual. This study examined whether there were past transitions in the ecological regime of Kings Billabong, a shallow River Murray wetland in southeast Australia. A 94-cm-long core, covering c. 90 years of age, was analysed at 1 cm resolution for subfossil cladocerans, diatoms and other proxies. Prior to river regulation (c. 1930), the littoral to planktonic ratios of cladocerans and diatoms, and bulk sediment δ¹³C values were high, while the period from c. 1930 to c. 1970 experienced considerable changes to the wetland ecosystem. The abrupt nature of changes of planktonic cladocerans and diatoms, particularly after the onset of river regulation (1930s), was triggered by inundation, high rates of sedimentation and shifts in bulk sediment δ¹⁵N values. However, the transition of a once littoral-dominated community, to one favouring an increasingly turbid, plankton-dominated trophic condition following river regulation was relatively slow and lasted for decades. The progression to a new regime was likely delayed by the partial recovery of submerged plant communities and related internal dynamics.     

湖泊生态恢复的基本原理与实现

当前我国湖泊污染及富营养化问题非常严重。湖泊治理的一个有效途径就是恢复水生植物,通过草型湖泊生态系统的培植来达到控制富营养化和净化水质的目的。但是,迄今为止,只有在局部水域或滨岸地区获得成功,恢复的水生植物主要是挺水植物或漂浮植物。鲜有全湖性的水生植物恢复和生态修复成功的例子。原因是对湖泊生态系统退化及其修复的机理了解甚少。实际上,环境条件不同决定了生态系统类型的不同,只有通过环境条件的改变才能实现生态系统的转变。利用草型湖泊生态系统来净化水质,其实质是利用生态系统对环境条件的反馈机制。但是,这种反馈无法从根本上改变其环境条件,因此其作用是有限的,不宜过分夸大。以往许多湖泊生态修复的工作之所以鲜有成功的例子,原因就是过于注重水生植物种植本身,而忽视了水生植物生长所需的环境条件的分析和改善。实施以水生植物恢复为核心的生态修复需要一定的前提条件。就富营养化湖泊生态恢复而言,这些环境条件包括氮磷浓度不能太高,富含有机质的沉积物应该去除,风浪不能太大以免对水生植物造成机械损伤,水深不能太深以免影响水生植物光合作用,鱼类种群结构应以食肉性鱼为主等等。因此,在湖泊污染很重或者氮磷负荷很高的情况下,寻求以沉水植物为核心的湖泊生态恢复来改善水质是不切实际的。为此,提出湖泊治理应该遵循先控源截污、后生态恢复,即先改善基础环境,后实施生态恢复的战略路线。     

Abrupt regime shifts in space and time along rivers and connected lake systems

Regime shifts between clear and turbid water states are commonly found in shallow lakes. These shifts are attributed to a positive feedback between water clarity and submerged macrophytes (underwater plants). Altering the retention time of the water may influence these interactions and thus potentially reduce the probability of alternative stable states. Here we assessed the effect of water retention time on the occurrence of alternative states in water quality of flushed lakes, chains of lakes and rivers using a spatially explicit simple model. Our results indicate that increased flushing of lakes rapidly decreases the range of parameters with alternative stable states up to their total disappearance at a flushing rate of about 50% the algal growth rate. Similarly, in a chain of lakes or in rivers with low flowing velocity, our model predicts that alternative stable states can only occur for systems with a high retention time. Despite the lack of hysteresis at lower water retention times, we predict that abrupt changes between clear and turbid states are still possible both in time and in space. Over a wide range of parameters, the equilibrium state of the chain of lakes shows a steep gradient of vegetation cover. Further, the transient dynamics of the model often include rapid shifts in time. For example, a local regime shift that occurs upstream may propagate through the whole lake chain or river due to a domino effect. All results of the simple model could qualitatively be reproduced with a more mechanistic model. The abrupt rather than gradual response of submerged macrophytes to reduced turbidity levels still makes river systems rather resilient to management measure. The importance of the initial turbidity and the observed domino effect suggest that restoration measures should start upstream and that these measures should eventually trigger regime shifts downstream.     

The pathway to noxious cyanobacteria blooms in lakes: the food web as the final turn

1. Cyanobacteria blooms have long been the focus of limnological research as they often represent the 'end member' of limnological deterioration under various human impacts. Research over the past several decades has greatly illuminated the ecological factors promoting cyanobacteria blooms but controversy and confusion surround the successful integration of this diverse body of work.
2. In this opinion article I attempt to integrate well-known aspects of cyanobacteria bloom ecology (such as the roles of nutrient loading, N:P ratio, and mixing conditions) with more recent developments that highlight the importance of feedbacks within the food web in regulating cyanobacteria blooms. Food-web feedbacks involving stoichiometric mechanisms appear to be particularly important, as accumulating data indicate that the food web influences cyanobacteria not just by regulating the rate of grazing mortality. Rather, trophic interactions may also regulate cyanobacteria dynamics by altering the consumer-driven nutrient recycling regime in a way that shifts the competitive advantage away from cyanobacteria.
3. Viewed in this way, cyanobacteria blooms can be seen as probabilistic events that are the end result of a series of key mechanisms involving nutrient loading, physical mixing conditions, and trophic interactions. To successfully manage lake water quality we should take advantage of each node of contingency leading to undesirable blooms. In doing so we will also have a more coherent scientific message to communicate with those directly involved with the socioeconomic politics of water quality decision making.     

Macrophytes moderate the taxonomic and functional composition of phytoplankton assemblages during a nutrient loading experiment

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基于管理目标的湖泊生态系统动力学

基于生态系统管理的目标,在对相关研究分析的基础上,依据生态系统生态学、淡水生态学的理论,提出了湖泊生态系统动力学研究的2个理论基础：生态系统管理和生态系统特征.在此基础上,分析得到湖泊生态系统动力学的研究方法体系,主要包括研究内容与技术路线、关键问题识别和动力学模拟、湖泊生态系统的适应性管理决策等部分.其中,湖泊生态系统结构和过程、湖泊中食物网营养动力学研究、生源要素循环、湖泊中关键过程的生态作用以及湖泊生态系统动力学模拟是研究的核心问题.此后,以P为主要的生源要素,将生态系统分为3个子过程：入流、出流和内部反馈,并以此建立了湖泊生态系统动力学的模型框架,以辅助于湖泊的生态系统管理.     

Effect of macrophyte community composition and nutrient enrichment on plant biomass and algal blooms

Submerged freshwater macrophytes decline with increasing eutrophication. This has consequences for ecosystem processes in shallow lakes and ponds as macrophytes can reduce algal blooms under eutrophic conditions. We hypothesize that the productivity of submerged vegetation, biomass change under eutrophication and the suppression of algal blooms may be affected by macrophyte community composition. To test our hypothesis, we established three macrophyte community types in 36 fishless experimental ponds: one dominated by the oligotrophic species Chara globularis, one dominated by the eutrophic species Potamogeton pectinatus and a diverse vegetation which became co-dominated by Elodea nuttallii and C. globularis, and we fertilized half of the ponds.The macrophyte communities produced different amounts of biomass and they responded differently to fertilization. The community dominated by Potamogeton produced the lowest overall biomass, but was not affected by nutrient addition. The communities dominated by Chara and co-dominated by Elodea and Chara produced more than four-fold the amount of biomass produced in Potamogeton communities under oligotrophic conditions, but were strongly negatively affected by nutrient addition.Phytoplankton abundance did not differ significantly among the plant community types, but showed large variation within community types. There was a significant negative relationship between spring macrophyte biomass and the probability of summer algal blooms. The occurrence of algal blooms coincided with low daphnid densities and high pH (>10).We conclude that the macrophyte community composition, characterized by the dominant species, strongly affected the amount of biomass production as well as the short-term response of the vegetation to nutrient enrichment. Macrophyte community composition had no direct effect on algal blooms, but can affect the occurrence of algal blooms indirectly as these occurred only in ponds with low (<100 g/m² DW) spring macrophyte biomass.     

Benthic algae compensate for phytoplankton losses in large aquatic ecosystems

Anthropogenic activities can induce major trophic shifts in aquatic systems, yet we have an incomplete understanding of the implication of such shifts on ecosystem function and on primary production (PP) in particular. In recent decades, phytoplankton biomass and production in the Laurentian Great Lakes have declined in response to reduced nutrient concentrations and invasive mussels. However, the increases in water clarity associated with declines in phytoplankton may have positive effects on benthic PP at the ecosystem scale. Have these lakes experienced oligotrophication (a reduction of algal production), or simply a shift in autotrophic structure with no net decline in PP? Benthic contributions to ecosystem PP are rarely measured in large aquatic systems, but our calculations based on productivity rates from the Great Lakes indicate that a significant proportion (up to one half, in Lake Huron) of their whole‐lake production may be benthic. The large declines (5–45%) in phytoplankton production in the Great Lakes from the 1970s to 2000s may be substantially compensated by benthic PP, which increased by up to 190%. Thus, the autotrophic productive capacity of large aquatic ecosystems may be relatively resilient to shifts in trophic status, due to a redirection of production to the near‐shore benthic zone, and large lakes may exhibit shifts in autotrophic structure analogous to the regime shifts seen in shallow lakes.     

Plant community structure determines primary productivity in shallow,eutrophic lakes

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Macrophyte refuges, prey behaviour and trophic interactions: consequences for lake water clarity

Macrophytes may enhance grazing on phytoplankton by providing a refuge for zooplankton against fish predation. Loss of macrophytes can trigger sudden degradation of water clarity (regime shift) in lakes. However, the presence of piscivores may drive planktivorous fish to take refuge amongst littoral macrophytes. To address the possibility of regime shifts, I here constructed an empirically based model that combined population dynamics of organisms with game theory for optimal habitat selection, taking into consideration the trophic structure, lake size and eutrophication. The model showed that macrophytes generally acted as a refuge for zooplankton, rather than for fish. The model predicted that regime shifts were more likely in small, shallow lakes and that the presence of macrophytes raised the possibility of regime shifts. The present study demonstrated that the fast dynamics of animal behaviour could lead to regime shifts, in connection with slower variables such as nutrient loading.     

浅水湖泊生态系统稳态转换的阈值判定方法

浅水湖泊生态系统对人类干扰的反应会随着干扰力度的改变或增强而出现突然的变化,即发生稳态转换;对其机理和驱动机制的揭示将有助于对湖泊富营养化的控制及恢复.基于“多稳态”理论的稳态转换研究已广泛开展,但对浅水湖泊生态系统稳态转换的驱动机制结论各异,采用的阈值判定方法相差很大,主要有实验观测、模型模拟和统计分析3种.实验观测多关注少数特定指标,指标筛选过程复杂且工作量大;模型模拟虽能从较为全面的尺度上理解生态系统稳态变化的特征和主要机理过程,但在模型误差和不确定性的处理等问题上尚存在不足;统计分析方法基于对长时间序列数据的统计变化规律分析,用以判断或者预警稳态转换现象的发生,是目前最为常用的方法.目前稳态转换领域的研究大都是对已发生的稳态转换进行机制分析或过程反演,对未来预测与预警的问题仍然亟需加强.     

POSITIVE FEEDBACK AND THE DEVELOPMENT AND PERSISTENCE OF ECOSYSTEM DISRUPTIVE ALGAL BLOOMS1

Harmful algal blooms (HABs) have occurred with increasing frequency in recent years with eutrophication and other anthropogenic alterations of coastal ecosystems. Many of these blooms severely alter or degrade ecosystem function, and are referred to here as ecosystem disruptive algal blooms (EDABs). These blooms are often caused by toxic or unpalatable species that decrease grazing rates by planktonic and benthic herbivores, and thereby disrupt the transfer of nutrients and energy to higher trophic levels, and decrease nutrient recycling. Many factors, such as nutrient availability and herbivore grazing have been proposed to separately influence EDAB dynamics, but interactions among these factors have rarely been considered. Here we discuss positive feedback interactions among nutrient availability, herbivore grazing, and nutrient regeneration, which have the potential to substantially influence the dynamics of EDAB events. The positive feedbacks result from a reduction of grazing rates on EDAB species caused by toxicity or unpalatability of these algae, which promotes the proliferation of the EDAB species. The decreased rates also lower grazer‐mediated recycling of nutrients and thereby decrease nutrient availability. Since many EDAB species are well‐adapted to nutrient‐stressed environments and many exhibit increased toxin production and toxicity under nutrient limitation, positive feedbacks are established which can greatly increase the rate of bloom development and the adverse effects on the ecosystem. An understanding of how these feedbacks interact with other regulating factors, such as benthic/pelagic nutrient coupling, physical forcing, and life cycles of EDAB species provides a substantial future challenge.     

Strong spatial differentiation of N and P deficiency,primary productivity and community composition between Nyanza Gulf and Lake Victoria (Kenya,East Africa) and the implications for nutrient management

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洱海蓝藻爆发的时空特征及影响因子

湖泊生态环境对区域气候变化和流域人类活动十分敏感。随着流域的持续开发,洱海作为云贵高原第二大淡水湖泊面临着严重的生态与环境问题,主要包括水质恶化和生态功能的衰退,其中蓝藻水华问题尤为突出。针对湖泊现代监测数据存在时间序列较短、连续监测记录缺乏、监测位点不完全一致等问题,应用沉积物记录开展色素等多指标分析和环境变化重建研究,并对洱海湖区南、中、北3个湖盆的沉积物记录进行对比分析,从而探讨洱海富营养化与蓝藻爆发的历史与变化特征,并识别藻类响应模式的空间异同。沉积物色素记录结果表明,洱海蓝藻生物量变化具有明显的时空差异性,呈现由南至北,先后增加、最后呈现蓝藻水华全湖性持续爆发的模式。进一步的简约模型方差分解结果表明气候变暖和营养盐富集是洱海蓝藻生物量变化的主要驱动因子,此外相对较浅的南部湖盆还受到水位波动、水动力减弱、水生植物演化的综合影响。因此,在气候变暖的背景下,控制水体营养盐输入、合理调控湖泊水位、提高水体透明度并恢复水生植物是控制洱海蓝藻水华爆发和进行生态恢复的重要措施。     

蓝藻堆积和螺类牧食对苦草生长的影响

设计了双因素四组处理(对照组,加螺组,加藻组,螺藻组)的室外受控实验,模拟湖泊沿岸带水华蓝藻的堆积以及底栖螺类的牧食活动对沉水植物苦草生长的影响。结果表明:蓝藻堆积(水体叶绿素a浓度为220μg/L)对苦草的生长具有明显的抑制作用,和对照组以及加螺组相比,加藻组和螺藻组中苦草的相对生长率分别下降了40.9%和36.4%,分株数也分别下降了56.4%和64.1%,分析认为蓝藻在水体表层堆积所产生的遮光可能是抑制底层苦草生长的主要原因。然而,环棱螺能在一定程度上促进苦草的生长,加螺组和螺藻组中苦草的相对生长率和分株数分别要明显高于对照组和加藻组,这可能要归因于螺类的牧食去除了沉水植物表面附着生物。实验中蓝藻堆积和螺类牧食对苦草的各项生长指标均无显著的交互作用,但蓝藻对苦草生长的抑制作用要远大于螺类对植物生长的促进作用。研究证实了在富营养浅水湖泊中,水华蓝藻在湖泊沿岸带的堆积会严重胁迫沉水植物的生长,而底栖螺类的牧食活动则能在一定程度上提高植物在不良环境下的生存能力。