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。     

Resource availability determines the importance of niche‐based versus stochastic community assembly in grasslands

Niche‐based selection and stochastic processes can operate simultaneously to generate spatial and temporal variation in species composition. Yet, the conditions under which ecological dynamics are dominated by niche‐based versus stochastic processes are poorly understood. Using a field experiment in early‐successional temperate grassland and null models of beta diversity, this study investigates the effects of soil nutrient supply on the relative importance of niche‐based selection versus stochastic dynamics for variation in species composition among sites. Nutrient availability was manipulated experimentally, individual seed mixtures with 25 species were sown in each experimental plot, and then stochastic and deterministic niche‐based assembly processes were allowed to happen. We found that compositional variation among grassland plots with low nutrient supply was driven by stochastic immigration and extinctions. In contrast, nutrient enrichment reduced the importance of stochasticity and imposed a deterministic environmental filter that homogenized communities through the selection of few species with greater competitive ability for light. This demonstrates that soil nutrient availability is a critical environmental feature that dictates the degree to which terrestrial plant communities are controlled by niche‐based selection versus stochastic assembly processes. Our study shows further that alternative states of eutrophic grasslands emerge from initial stochastic variation in the composition of a particular functional group of species that can become dominant at high nutrient supply. We discuss potential mechanisms underlying the shift from stochastic to niche‐driven dynamics along soil nutrient gradients.     

On evolution under asymmetric competition

The evolutionary consequences of asymmetric competition between species are poorly understood in comparison with symmetric competition. A model for evolution of body size under asymmetric competition within and between species is described. The model links processes operating at the scale of the individual to that of macroscopic evolution through a stochastic mutation–selection process. Phase portraits of evolution in a phenotype space characteristically show character convergence and parallel character shifts, with character divergence being relatively uncommon. The asymptotic states of evolution depend very much on the properties of asymmetric competition. Given relatively weak asymmetries between species, a single equilibrium point exists; this is a local attractor, and its position is determined by the intra- and interspecific asymmetries. When the asymmetries are made stronger, several fixed points may come about, creating further equilibrium points which are local attractors. It is also possible for periodic attractors to occur; such attractors comprise Red Queen dynamics with phenotype values that continue to change without ever settling down to constant values. From certain initial conditions, evolution leading to extinction of one of the species is also a likely outcome.     

Shallow lakes theory revisited: various alternative regimes driven by climate,nutrients, depth and lake size

Shallow lakes have become the archetypical example of ecosystems with alternative stable states. However, since the early conception of that theory, the image of ecosystem stability has been elaborated for shallow lakes far beyond the simple original model. After discussing how spatial heterogeneity and fluctuation of environmental conditions may affect the stability of lakes, we review work demonstrating that the critical nutrient level for lakes to become turbid is higher for smaller lakes, and seems likely to be affected by climatic change too. We then show how the image of just two contrasting states has been elaborated. Different groups of primary producers may dominate shallow lakes, and such states dominated by a particular group may often represent alternative stable states. In tropical lakes, or small stagnant temperate waters, free-floating plants may represent an alternative stable state. Temperate shallow lakes may be dominated alternatively by charophytes, submerged angiosperms, green algae or cyanobacteria. The change of the lake communities along a gradient of eutrophication may therefore be seen as a continuum in which gradual species replacements are interrupted at critical points by more dramatic shifts to a contrasting alternative regime dominated by different species. The originally identified shift between a clear and a turbid state remains one of the more dramatic examples, but is surely not the only discontinuity that can be observed in the response of these ecosystems to environmental change.     

Seasonal dynamics of macrophytes and phytoplankton in shallow lakes: a eutrophication‐driven pathway from plants to plankton?

1. Seasonal relationships between macrophyte and phytoplankton populations may alter considerably as lakes undergo eutrophication. Understanding of these changes may be key to the interpretation of ecological processes operating over longer (decadal‐centennial) timescales. 2. We explore the seasonal dynamics of macrophytes (measured twice in June and August) and phytoplankton (measured monthly May–September) populations in 39 shallow lakes (29 in the U.K. and 10 in Denmark) covering broad gradients for nutrients and plant abundance. 3. Three site groups were identified based on macrophyte seasonality; 16 lakes where macrophyte abundance was perennially low and the water generally turbid (‘turbid lakes’); 7 where macrophyte abundance was high in June but low in August (‘crashing’ lakes); and 12 where macrophyte abundance was high in both June and August (‘stable’ lakes). The seasonal behaviour of the crashing and turbid lakes was extremely similar with a consistent increase in nutrient concentrations and chlorophyll‐a over May–September. By contrast in the stable lakes, seasonal changes were dampened with chlorophyll‐a consistently low (<10–15 μg L^?1) over the entire summer. The crashing lakes were dominated by one or a combination of Potamogeton pusillus, Potamogeton pectinatus and Zannichellia palustris, whereas Ceratophyllum demersum and Chara spp. were more abundant in the stable lakes. 4. A long‐term loss of macrophyte species diversity has occurred in many shallow lakes affected by eutrophication. One common pathway is from a species‐rich plant community with charophytes to a species‐poor community dominated by P. pusillus, P. pectinatus and Z. palustris. Such compositional changes may often be accompanied by a substantial reduction in the seasonal duration of plant dominance and a greater tendency for incursions by phytoplankton. We hypothesise a slow‐enacting (10–100 s years) feedback loop in nutrient‐enriched shallow lakes whereby increases in algal abundance are associated with losses of macrophyte species and hence different plant seasonal strategies. In turn such changes may favour increased phytoplankton production thus placing further pressure on remaining macrophytes. This study blurs the distinction between so‐called turbid phytoplankton‐dominated and clear plant‐dominated shallow lakes and suggests that plant loss from them may be a gradual process.     

Wetland macrophyte diversity, trophic status and phytoplankton： a case study of the cascading effects of an invasive herbivore

Shallow water bodies can exist in alternative stable states, a clear water state with high coverage of macrophytes or a turbid state with high phytoplankton biomass. The alternative equilibria hypothesis has been proposed to explain the occurrence of the alternative stable states (Scheffer et al., 1993)[1], which assumes that: 1),     

Mathematical modeling of colony formation in algal blooms: phenotypic plasticity in cyanobacteria

In this paper, we analyzed a mathematical model of algal-grazer dynamics, including the effect of colony formation, which is an example of phenotypic plasticity. The model consists of three variables, which correspond to the biomasses of unicellular algae, colonial algae, and herbivorous zooplankton. Among these organisms, colonial algae are the main components of algal blooms. This aquatic system has two stable attractors, which can be identified as a zooplankton-dominated (ZD) state and an algal-dominated (AD) state, respectively. Assuming that the handling time of zooplankton on colonial algae increases with the colonial algae biomass, we discovered that bistability can occur within the model system. The applicability of alternative stable states in algae-grazer dynamics as a framework for explaining the algal blooms in real lake ecosystems, thus, seems to depend on whether the assumption mentioned above is met in natural circumstances.     

Catastrophic Regime Shift in Water Reservoirs and S?o Paulo Water Supply Crisis

The relation between rainfall and water accumulated in reservoirs comprises nonlinear feedbacks. Here we show that they may generate alternative equilibrium regimes, one of high water-volume, the other of low water-volume. Reservoirs can be seen as socio-environmental systems at risk of regime shifts, characteristic of tipping point transitions. We analyze data from stored water, rainfall, and water inflow and outflow in the main reservoir serving the metropolitan area of São Paulo, Brazil, by means of indicators of critical regime shifts, and find a strong signal of a transition. We furthermore build a mathematical model that gives a mechanistic view of the dynamics and demonstrates that alternative stable states are an expected property of water reservoirs. We also build a stochastic version of this model that fits well to the data. These results highlight the broader aspect that reservoir management must account for their intrinsic bistability, and should benefit from dynamical systems theory. Our case study illustrates the catastrophic consequences of failing to do so.     

Migratory benthic fishes may induce regime shifts in a tropical floodplain pond

1. Alternative states are a widely recorded phenomenon in shallow lakes, which may shift between turbid‐ and clear‐water conditions. Here, we investigate whether such shifts in a tropical floodplain pond may be related to the effect of the flood pulse regime on the community structures of fish and macrophytes. 2. Using a long‐term data set, we demonstrate how benthic fish migration together with colonisation by submerged plants affected the transition from a turbid to a macrophyte‐dominated state in a floodplain pond without top‐down control. 3. In our study, the turbid state occurred mostly during low water phases and was largely characterised by high values for the biomass of benthic fish, chlorophyll‐a and total phosphorous. 4. During the period of rising water levels, the migration of benthic fish out of the pond occurs simultaneously with the establishment of submerged plants, while water turbidity decreases along with phytoplankton and nutrient concentrations, inducing a clear‐water phase. However, when submerged plants are absent and fish migration is low, a transient state is generated. 5. We suggest that, in contrast to temperate ponds and shallow lakes, where the main driving mechanisms establishing alternative states are related to cascading effects via the food chain, in tropical ponds and shallow lakes it is resuspension of sediments by benthic fish that plays the most significant role in establishing alternative states. However, the effect of the flood pulse regime plays an important role in the temporal dynamics of fish community structure by controlling benthic fish migration.     

Positive interactions, discontinuous transitions and species coexistence in plant communities

The population and community level consequences of positive interactions between plants remain poorly explored. In this study we incorporate positive resource-mediated interactions in classic resource competition theory and investigate the main consequences for plant population dynamics and species coexistence. We focus on plant communities for which water infiltration rates exhibit positive dependency on plant biomass and where plant responses can be improved by shading, particularly under water limiting conditions. We show that the effects of these two resource-mediated positive interactions are similar and additive. We predict that positive interactions shift the transition points between different species compositions along environmental gradients and that realized niche widths will expand or shrink. Furthermore, continuous transitions between different community compositions can become discontinuous and bistability or tristability can occur. Moreover, increased infiltration rates may give rise to a new potential coexistence mechanism that we call controlled facilitation.     

Potential for Sudden Shifts in Transient Systems: Distinguishing Between Local and Landscape-Scale Processes

Thorough understanding of the potential for threshold dynamics and catastrophic shifts to occur in natural systems is of great importance for ecosystem conservation and restoration. However, verifying the presence of alternative stable states, one of the theoretical explanations for sudden shifts in natural systems, has proven to be a major challenge. We examine processes on local and landscape scales in salt-marsh pioneer zones, to assess the presence of alternative stable states in this system. To that end, we investigated the presence of typical characteristics of alternative stable states: bimodality and threshold dynamics. We also studied whether vegetation patches remained stable over long time periods. Analysis of false-color aerial photographs revealed clear bimodality in plant biomass distribution. By transplanting Spartina anglica plants of three different biomass classes on three geographically different marshes, we showed that a biomass threshold limits the establishment of Spartina patches, potentially explaining their patchy distribution. The presence of bimodality and biomass thresholds points to the presence of alternative stable states and the potential for sudden shifts, at small, within-patch scales and on short time scales. However, overlay analysis of aerial photographs from a salt marsh in The Netherlands, covering a time span of 22 years, revealed that there was little long-term stability of patches, as vegetation cover in this area is slowly increasing. Our results suggest that the concept of alternative stable states is applicable to the salt-marsh pioneer vegetation on small spatio-temporal scales. However, the concept does not apply to long-term dynamics of decades or centuries of heterogeneous salt-marsh pioneer zones, as landscape-scale processes may determine the large-scale dynamics of salt marshes. Hence, our results provide the interesting perspective that threshold dynamics may occur in systems with, on the long term, only a single stable state. Author Contributions: Bregje van Wesenbeeck executed the major part of this research and wrote this paper. Johan van de Koppel was involved in every part of this study and restructured major parts of this paper. Peter Herman helped with statistical analyses. Mark Bertness adjusted the design of this study and provided new views. Daphne van der Wal performed GIS analyses. Jan Bakker supervised and facilitated field work. Tjeerd Bouma supervised design of study and paper and substantially contributed to the writing of this paper. All authors commented on several drafts of this paper.     

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.     

Climate change causes critical transitions and irreversible alterations of mountain forests

Mountain forests are at particular risk of climate change impacts due to their temperature limitation and high exposure to warming. At the same time, their complex topography may help to buffer the effects of climate change and create climate refugia. Whether climate change can lead to critical transitions of mountain forest ecosystems and whether such transitions are reversible remain incompletely understood. We investigated the resilience of forest composition and size structure to climate change, focusing on a mountain forest landscape in the Eastern Alps. Using the individual‐based forest landscape model iLand, we simulated ecosystem responses to a wide range of climatic changes (up to a 6°C increase in mean annual temperature and a 30% reduction in mean annual precipitation), testing for tipping points in vegetation size structure and composition under different topography scenarios. We found that at warming levels above +2°C a threshold was crossed, with the system tipping into an alternative state. The system shifted from a conifer‐dominated landscape characterized by large trees to a landscape dominated by smaller, predominantly broadleaved trees. Topographic complexity moderated climate change impacts, smoothing and delaying the transitions between alternative vegetation states. We subsequently reversed the simulated climate forcing to assess the ability of the landscape to recover from climate change impacts. The forest landscape showed hysteresis, particularly in scenarios with lower precipitation. At the same mean annual temperature, equilibrium vegetation size structure and species composition differed between warming and cooling trajectories. Here we show that even moderate warming corresponding to current policy targets could result in critical transitions of forest ecosystems and highlight the importance of topographic complexity as a buffering agent. Furthermore, our results show that overshooting ambitious climate mitigation targets could be dangerous, as ecological impacts can be irreversible at millennial time scales once a tipping point has been crossed.     

Forecasting Bifurcations from Large Perturbation Recoveries in Feedback Ecosystems

Forecasting bifurcations such as critical transitions is an active research area of relevance to the management and preservation of ecological systems. In particular, anticipating the distance to critical transitions remains a challenge, together with predicting the state of the system after these transitions are breached. In this work, a new model-less method is presented that addresses both these issues based on monitoring recoveries from large perturbations. The approach uses data from recoveries of the system from at least two separate parameter values before the critical point, to predict both the bifurcation and the post-bifurcation dynamics. The proposed method is demonstrated, and its performance evaluated under different levels of measurement noise, with two ecological models that have been used extensively in previous studies of tipping points and alternative steady states. The first one considers the dynamics of vegetation under grazing; the second, those of macrophyte and phytoplankton in shallow lakes. Applications of the method to more complex situations are discussed together with the kinds of empirical data needed for its implementation.     

On evolution under symmetric and asymmetric competitions

This paper considers the coevolution of phenotypic traits in a community comprising two competitive species subject to strong Allee effects. Firstly, we investigate the ecological and evolutionary conditions that allow for continuously stable strategy under symmetric competition. Secondly, we find that evolutionary suicide is impossible when the two species undergo symmetric competition, however, evolutionary suicide can occur in an asymmetric competition model with strong Allee effects. Thirdly, it is found that evolutionary bistability is a likely outcome of the process under both symmetric and asymmetric competitions, which depends on the properties of symmetric and asymmetric competitions. Fourthly, under asymmetric competition, we find that evolutionary cycle is a likely outcome of the process, which depends on the properties of both intraspecific and interspecific competition. When interspecific and intraspecific asymmetries vary continuously, we also find that the evolutionary dynamics may admit a stable equilibrium and two limit cycles or two stable equilibria separated by an unstable limit cycle or a stable equilibrium and a stable limit cycle.     

The effect of evolution on host-parasitoid systems

It is well known that a simple first-order difference equation can exhibit complex population dynamics, such as sustained oscillations and chaos. An interesting problem is whether such oscillatory dynamics are expected to occur in real populations. This paper assumes that the resident system is composed of 1-host and 1-parasitoid and that only the host is allowed to evolve, but not the parasitoid. Based on the invasibility of a host to host-parasitoid systems, we investigate the dynamics of the host-parasitoid system favored by natural selection. We consider two cases. In the first case, the host's evolution involving both the intrinsic growth rate and the sensitivity to density is considered. In the second case, the host's evolution involving both the intrinsic growth rate and the vulnerability to the parasitoid is considered. In both cases, we see that the dynamics with a stable equilibrium will not be favored by natural selection without the trade-off between the host's traits which are allowed to evolve. The host-parasitoid system with a stable equilibrium will be eventually invaded by a host type that develops an unstable equilibrium with the parasitoid. If there is a trade-off between the host's traits which are allowed to evolve, a host-parasitoid system with a stable equilibrium can be favored by natural selection.     

Competition and evolution along environmental gradients: patterns, boundaries and sympatric divergence

The present study examined how competitive interactions and environmental conditions generate species boundaries and determine species distributions. A spatially explicit, quantitative genetic, two-species competition model was used to manipulate the strengths of competition, gene flow and local adaptation along environmental gradients. This allowed us to assess the long-term persistence of each species and whether the ranges they inhabited had boundaries in space or were unlimited. We found that a species boundary arises along less steep environmental gradients when the strength of stabilizing selection and diversifying selection are similar. We also found that a species boundary may arise along shallow environmental gradients if interspecific competition is more intense than intraspecific, which relaxes previous requirements for steep gradients for generating range limits. We determined an analytical form for the critical environmental gradient as a function of ecological and genetic parameters at which a species boundary is expected to arise by competition. Results suggest an alternative to resource competition as an explanation for phenotypic divergence between sympatric competitors. Competitors sharing a trait that is under stabilizing selection along an environmental gradient may segregate spatially and evolve in different regions, with phenotypic sympatric divergence reflecting the resulting clines. Along various types of environmental gradients, variation in stabilizing selection intensities could lead to contrasting patterns in the distribution of species. For stabilizing selection strengths in accord with field data estimates, this study predicts that the level of sympatric character divergence would be limited along environmental gradients.     

Waterfowl,macrophytes, and the clear water state of shallow lakes

The importance of lake ecosystems for waterfowl remains a topic of debate. In order to assess how temporal variations in lake features, specifically shifts between alternative stable states, may interact with the waterfowl fauna, we performed a long-term (22 years) study of the shallow Lake Krankesjön, southern Sweden. Lower total numbers of waterfowl occurred during periods with low macrophyte cover and turbid water, than when submersed macrophytes flourished and the water was clear. Some specific functional groups of waterfowl, such as herbivores, invertebrate, and fish feeders, showed a positive relation to clear water and high macrophyte cover. Hence, our data suggest that some migratory waterfowl may select lakes based on water quality, thereby adjusting their large-scale migratory routes. On the other hand, omnivorous waterfowl exhibited their highest abundances during turbid conditions. Furthermore, waterfowl not primarily relying on food from the lake showed no response to fluctuations in turbidity or macrophyte cover, but followed regional trends in population dynamics. In our study lake, L. Krankesjön, we estimated that waterfowl remove less than 3% of the macrophyte biomass during a stable clear-water state with lush macrophyte beds. However, during transition periods between alternative stable states, when macrophyte biomass is lower and the plants already stressed, the consumption rate of waterfowl may have a stronger effect on lake ecosystem functioning.     

What evidence exists for alternative ecological regimes in salinising wetlands?

1. Land clearing in Australia's southwest has led to widespread salinisation of aquatic ecosystems. Four different ecological regimes (clear, submerged macrophyte‐dominated; clear, benthic microbial community‐dominated; turbid, phytoplankton‐dominated; and turbid, sediment‐dominated) have previously been identified in the salinising wetlands of this region. 2. Monitoring data from seven saline wetlands over an 18‐month period were used to evaluate whether a continuum, simple threshold or alternative regimes conceptual model (sensu Hydrobiologia, 200/201, 1990, 367; Hydrobiologia, 200/201, 1990, 475) most appropriately represented transitions between these ecological regimes. We also aimed to identify whether factors other than salinity played a major role in defining ecological regimes or causing shifts between them. 3. Ordination of biological variables revealed a separation of benthic microbial community‐dominated from submerged macrophyte‐dominated sites and times. The mean salinities of these two groups were very similar, suggesting that a salinity threshold was not responsible for benthic microbial versus macrophyte dominance. No other environmental variable was found to have a strong, direct influence on the groupings. 4. The dynamics of regime change in saline wetlands appear not to be driven by any single variable, but by the combined effects of salinity and water regime on species life histories and competitive abilities. Macrophytes were powerful competitors, able to germinate and establish under a range of salinities, turbidities and water depths, and were favoured by seasonal drying. 5. Data from the seven wetlands indicated that the continuum, simple threshold and alternative regimes conceptual models did not appropriately represent transitions between ecological regimes in seasonally drying wetlands. Macrophyte and benthic microbial regimes occurred at overlapping salinity levels, excluding both the continuum and threshold models, and the regular occurrence of drying appeared to preclude the alternative regimes model. Drying prevented the development of strong positive feedback mechanisms, which might otherwise have maintained the benthic microbial community‐dominated regime. We hypothesise that an alternative regimes model might still be valid for salinising ecosystems holding permanent water.     

Effects of plant evolution on nutrient cycling couple aboveground and belowground processes

Plant strategies for nutrient acquisition and recycling are key components of ecosystem functioning. How the evolution of such strategies modifies ecosystem functioning and services is still not well understood. In the present work, we aim at understanding how the evolution of different phenotypic traits link aboveground and belowground processes, thereby affecting the functioning of the ecosystem at different scales and in different realms. Using a simple model, we follow the dynamics of a limiting nutrient inside an ecosystem. Considering trade-offs between aboveground and belowground functional traits, we study the effects of the evolution of such strategies on ecosystem properties (amount of mineral nutrient, total plant biomass, dead organic matter, and primary productivity) and whether such properties are maximized. Our results show that when evolution leads to a stable outcome, it minimizes the quantity of nutrient available (following Tilman’s R* rule). We also show that considering the evolution of aboveground and belowground functional traits simultaneously, total plant biomass and primary productivity are not necessarily maximized through evolution. The coupling of aboveground and belowground processes through evolution may largely diminish predicted standing biomass and productivity (extinction may even occur) and impact the evolutionary resilience (i.e., the return time to previous phenotypic states) of the ecosystem in the face of external disturbances. We show that changes in plant biomass and their effects on evolutionary change can be understood by accounting for the links between nutrient uptake and mineralization, and for indirect effects of nutrient uptake on the amount of detritus in the system.