Evolutionary emergence of alternative stable states in shallow lakes

Ecosystems under stress may respond abruptly and irreversibly through tipping points. Although mechanisms leading to alternative stable states are much studied, little is known about how such ecosystems could have emerged in the first place. We investigate whether evolution by natural selection along resource gradients leads to bistability, using shallow lakes as an example. There, tipping points occur between two alternative states dominated by either submersed or floating macrophytes depending on nutrient loading. We model the evolution of macrophyte depth in the lake, identify the conditions under which the ancestor population diversifies and investigate whether alternative stable states dominated by different macrophyte phenotypes occur. We find that eco-evolutionary dynamics may lead to alternative stable states, but under restrictive conditions. Such dynamics require sufficient asymmetries in the acquisition of both light and nutrient. Our analysis suggests that competitive asymmetries along opposing resource gradients may allow bistability to emerge by natural selection.     

Submerged vegetation development in two shallow, eutrophic lakes

Submerged macrophyte vegetation in two shallow lakes in the Netherlands, Lake Veluwemeer and Lake Wolderwijd, has been affected by eutrophication in the late 1960's and 1970's. Recent changes in the vegetation occurred in the period following lake restoration measures. Between 1987 and 1993, the dominance of Potamogeton pectinatus decreased, while Charophyte meadows expanded over the same time interval. The pattern of change of the dominant macrophyte species might result from changes in the underwater light climate. Seasonally persistent clear water patches associated with the Chara meadows have been observed in the last few years. The interaction between submerged macrophyte vegetation succession and water transparency in the lakes is discussed.     

Alternative stable states in eutrophic,shallow freshwater systems: A minimal model

A simple mathematical model was constructed, describing the relationships between pike, bream, aquatic macrophytes and the nutrient loading of shallow lakes. The model is analyzed with the use of zero-isoclines. It is concluded that, over a certain range of nutrient concentrations, the ecological relations incorporated in the model can give rise to the existence of two alternative stable equilibria;viz. a turbid bream-dominated one, and a clear state in which pike and aquatic vegetation are abundant. Under oligotrophic conditions, the clear-water state represents the only stable equilibrium; however, at high nutrient levels, the clear state is absent, or only locally stable. The response of the model to both increase and decrease of the nutrient level is characterised by hysteresis. The results indicate that manipulation of fish densities as a measure to improve water quality is only likely to produce long-term results when the nutrient level is below a certain threshold.     

Alternative states in shallow lakes

Ecology of Shallow Lakes by Marten Scheffer Chapman and Hall (Population and Community Biology Series 22), 1998. ￡45.00/$79.65 hbk (xx+357 pages) ISBN 0 412 74920 3.     

Benthic carbon is inefficiently transferred in the food webs of two eutrophic shallow lakes

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Forms and distribution of inorganic phosphorus in sediments of two shallow eutrophic lakes in Florida

Physicochemical attributes were measured and aquatic macroinvertebrates were collected from six wetlands near Perth, Western Australia at three weekly intervals over a 13 month period from August 1988 to September 1989. The six wetlands encompassed a range of depths, pH, concuctivities, nutrient concentrations and colours. Temporal changes in the macroinvertebrate communities appeared to be related to seasonal changes in the physical and chemical characteristics of the wetlands. Community composition differed more between the less enriched wetlands then the higly enriched wetlands where communities were generally similar. High species richness was associated with seasonal drying. High macro invertebrate abundance appeared to be associated with the presence of either green algal or cyanobacterial blooms in the enriched wetlands. The highest macroinvertebrate biomass was recorded in wetlands with both cyanobacterial blooms and abundant macrophytes present.     

Mechanisms regulating abundance of submerged vegetation in shallow eutrophic lakes

Shallow eutrophic lakes tend to be either in a turbid state dominated by phytoplankton or in a clear-water state dominated by submerged macrovegetation. Recent studies suggest that the low water turbidity in the clear-water state is maintained through direct and indirect effects of the submerged vegetation. This study examined what mechanisms may cause a recession of the submerged vegetation in the clear-water state, and thereby a switch to the turbid state. The spatial distribution of submerged vegetation biomass was investigated in two shallow eutrophic lakes in the clear-water state in southern Sweden. Biomass of submerged vegetation was positively correlated with water depth and wave exposure, which also were mutually correlated, suggesting that mechanisms hampering submerged vegetation were strongest at shallow and/or sheltered locations. The growth of Myriophyllum spicatum, planted in the same substrate and at the same water depth, was compared between sheltered and wave exposed sites in two lakes. After 6 weeks the plants were significantly smaller at the sheltered sites, where periphyton production was about 5 times higher than at the exposed sites. Exclosure experiments were conducted to evaluate the effects of waterfowl grazing on macrophyte biomass. Potamogeton pectinatus growth was decreased by grazing, whereas M. spicatum was not affected. The effects were greater at a sheltered than at a wave-exposed site, and also negatively related to distance from the reed belt. These results suggest that competition from epiphytes and waterfowl grazing hamper the development of submerged vegetation at sheltered and/or shallow locations. An increased strength of these mechanisms may cause a recession of submerged vegetation in shallow eutrophic lakes in the clear-water state and thereby a switch to the turbid state. Received: 24 June 1996 / Accepted: 8 September 1996     

Ice cover and thaw events influence nitrogen partitioning and concentration in two shallow eutrophic lakes

Biogeochemistry - The frequency and duration of lake ice cover is rapidly declining in the Northern Hemisphere. Limited research in oligotrophic and mesotrophic lakes suggests that extended periods...     

大型浅水富营养化湖泊中蓝藻水华形成机理的思考

湖泊富营养化依然是我国目前以及今后相当长一段时期内的重大水环境问题。研究蓝藻水华的形成机制 ,对于科学预测湖泊中蓝藻水华的产生 ,并采取相应措施减少其带来的影响具有重要的生态和环境意义。为探索富营养化湖泊中蓝藻水华形成机理 ,综述了目前对我国大型浅水湖泊蓝藻水华成因研究现状和对水华形成机理的一般认识。分析了导致蓝藻水华形成的化学、物理和生物等主要环境因素 ,论述了蓝藻 ,尤其是微囊藻成为水华优势种的可能原因。认为对水华的形成需要全面认识 ,营养盐浓度的升高可能仅是蓝藻水华形成、且人们可以加以控制的因素之一 ;在探索水华成因时 ,不能仅仅局限于夏季蓝藻水华发生时环境特征的研究与观察 ,而应该提前关注蓝藻的越冬生理生态特征、春季复苏的生态诱导因子及其阈值以及在复苏后 ,蓝藻如何在生长过程中形成群体 ,并逐步成为湖泊水生生态系统中的优势种乃至形成水华的过程。并需要对蓝藻越冬的生存对策、蓝藻群体的形成的条件、蓝藻在春季复苏的触发条件及其生态阈值、以及蓝藻在与其它藻类种群竞争中取胜的生理生化特征有足够的认识。蓝藻水华的“暴发”是表观现象 ,其前提还是藻类一定的生物量 ,且是一个逐渐形成的过程。根据生态学的基本理论和野外对水华形成过程的原位观测     

Persistent internal phosphorus loading during summer in shallow eutrophic lakes

Nutrient availability, in particular of phosphorus (P), is a key factor for the structure and functioning of shallow lakes, and not least the sediment plays an important role by acting as both a nutrient source and sink. We used 21 years of monthly mass balance and lake water data from six shallow (mean depth = 1.2–2.7 m) and fast flushed (mean hydraulic retention time = 0.6–2.6 months) eutrophic Danish lakes (mean summer P concentrations ranging from 0.09 to 0.61 mg/l) to investigate long-term trends in yearly and seasonal patterns of P retention. To one of the lakes, the external P input was reduced by 70% in the early 1990s, whereas none of the other lakes have experienced major changes in external P loading for more than 20 years. All lakes showed a distinct seasonal pattern with high P concentrations and typically negative P retention during summer (up to ?300% of the external loading from May to August). During winter, P retention was overall positive (up to 50% of the external loading from December to April). Internal P loading from the sediment delayed lake recovery by approximately 10 years in the lake with the most recently reduced external loading, but in all the lakes net release of P from the sediment occurred during summer. P release in the six lakes has not abated during the past decade, indicating that the sediment of eutrophic and turbid shallow lakes remains a net source of P during summer. The seasonal variations in P retention became more pronounced with increasing P levels, and retention decreased with increasing temperature, but increased if clear water conditions were established.     

Indirect effects of fish community structure on submerged vegetation in shallow,eutrophic lakes: an alternative mechanism

The loss of submerged macrophytes during eutrophication of shallow lakes is a commonly observed phenomenon. The proximate reason for this decline is a reduction of available light due to increasing phytoplankton and/or epiphyton biomass. Here we argue that the ultimate cause for the transition from a macrophyte-dominated state to a phytoplankton-dominated state is a change in fish community structure. A catastrophic disturbance event (e.g. winterkill) acting selectively on piscivores, cascades down food chains, eventually reducing macrophyte growth through shading by epiphyton, an effect that is reinforced by increasing phytoplankton biomass. The transition back from the phytoplankton to the macrophyte state depends on an increase in piscivore standing stock and a reduction of planktivores. A conceptual model of these mechanisms is presented and supported by literature data and preliminary observations from a field experiment.     

Plant community structure determines primary productivity in shallow,eutrophic lakes

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Epiphyte biomass and elemental composition on submersed macrophytes in shallow eutrophic lakes

Hydrobiologia - Epiphytes cause a strong light attenuation for submersed macrophytes. Various abiotic and biotic factors determine epiphyte density, among them nutrients, water clarity and possibly...     

Effect of temperature on phosphorus sorption to sediments from shallow eutrophic lakes

The availability of phosphorus (P) in lakes is dependent on the sorption characteristics of the underlying sediments. Temperature is a crucial factor affecting the P sorption in sediments. The objective of this study was to evaluate the effect of temperature on sorption of P by sediments from two eutrophic lakes. The study was carried out using short-term batch experiments at 4, 20 and 30 °C. Phosphorus sorption kinetics, isotherms, fractionation and desorption were investigated. The P sorption was dependent on sediment type and temperature (p < 0.001). The Mei sediments showed a higher sorption rate and sorption capacity than Hua sediments. The P sorption kinetics were best described by a pseudo second order model (R² > 0.97). Activation energies derived from the kinetics rate constant indicated that P sorption onto the two sediments was controlled by a diffusion process. For both sediments, Freundlich model fit the P sorption isotherms well and the calculated apparent sorption heat was 6.37 kJ mol⁻¹ for Mei sediments and 8.67 kJ mol⁻¹ for Hua sediments. This indicated that P sorption onto both sediments was endothermic. Adding P significantly increased the soluble and loosely bound P (S/L-P), aluminum-bound P (Al-P) and iron-bound P (Fe-P) (p < 0.05). The amount of Al-P and Fe-P was markedly higher at 30 °C than at 4 °C (p < 0.05). Subsequent P desorption indicated that adsorbed P was highly labile, in particular for Hua sediment. The degree of P mobility that occurred during sediment sorption was inversely related to the temperature at the time of sorption. A significant relationship (R² = 0.978) between phosphorus sorption maximum and oxalate-extractable Fe and Al at different temperatures reflects that the amorphous contents of Fe and Al are responsible for the temperature effect on P sorption.     

Bacterial activities in shallow lakes – a comparison between extremely acidic and alkaline eutrophic hard water lakes

Hydrobiologia - Bacterial abundance and activity were investigated by DAPI-staining and 3H-thymidine incorporation method in samples from a very shallow, hypertrophic hard water lake and from...     

Cladoceran and rotifer grazing on bacteria and phytoplankton in two shallow eutrophic lakes: in situ measurement with fluorescent microspheres

Metazooplankton grazing on bacteria and on the phytoplanktonof various sizes was estimated in shallow eutrophic lakes Kaiavereand Võrtsjärv (Estonia) by in situ feeding experimentswith fluorescent microspheres (diameters 0.5 µm for bacteriaand 3, 6 and 24 µm for phytoplankton). Zooplankton communitycomposition, abundance and food density were important factorsdetermining grazing rates in these lakes. Cladocerans and rotifersfiltering rates (FR) and ingestion rates (IR) on bacteria andphytoplankton were several times higher in Lake Kaiavere wherebacterivorous rotifers and Daphnia contributed more to zooplanktonassemblage. While cladocerans were generally the main phytoplanktonconsumers, both lakes differed with respect to the groups ofbacterivores. Based on consumption of fluorescent microspheres,the metazooplankton grazing rates were relatively low and hadlow impact on production and standing stock of bacteria andingestible phytoplankton (<30 µm). On average, 0.5and 0.1% of standing stock of bacteria and 2.6 and 1.0% of standingstock of ingestible phytoplankton was grazed daily by metazooplanktonin lakes Kaiavere and Võrtsjärv, respectively. Thatcorresponded to daily grazing of 4.1% of the bacterial productionand 0.43% of the total primary production (PP) by metazooplanktonin Lake Kaiavere compared with 4.3 and 0.06% in Lake Võrtsjärv,respectively. The results suggest that the majority of consumptionof the bacterial and phytoplankton PP is most likely channelledthrough the microbial loop.     

A mosaic community of macrophytes for the ecological remediation of eutrophic shallow lakes

The restoration of macrophytes in eutrophic shallow lakes has been the focus of active research in the past decade. The approach of building up a mosaic community of macrophytes (MCMs) to improve water quality and remediate the ecosystems of lakes is proposed in this paper. Several species of floating, floating-leaved, and submerged macrophytes were introduced in experimental enclosures in eutrophic shallow lakes. These macrophytes were intercropped in small patches and formed mosaic communities of spatial and temporal combinations (spatial and seasonal mosaic patterns) in the lakes. Macrophytes can improve water transparency quickly and the MCM system can stabilize this clear water state over a long time in turbid eutrophic shallow lakes. The constructed MCM created heterogeneous habitats that are favourable for different macrophytes and for the growth and succession of other organisms, as well as for removing water pollutants. When the eutrophic water flowed through the MCM system at a retention time of 7 days, the removal efficiency rates of the MCM system for algae biomass, NH₄⁺–N, TN, TP and PO₄^3?–P were 58%, 66%, 60%, 72% and 80%, respectively.     

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...     

Detection of alternative stable states in marine communities

During the last 20 years there have been several attempts to test the theory of alternative stable states using marine systems. The results have been mixed, and there have been conflicting interpretations of the outcomes. In an attempt to resolve some of the problems, the theory and evidence for alternative stable states are reviewed. There are several different views of what is meant by alternative stable states, and so the key elements of the theory of alternative stable states are set in the context of marine systems. Appropriate experimental designs for detecting alternative states are discussed, and recent experimental studies in coral reefs, rocky intertidal systems, soft-sediment beds, and subtidal systems are briefly reviewed. Suggestions for improving experimental designs are proposed and unresolved issues are highlighted.