生态系统状态跃变的早期预警信号及检测

生态系统在气候变化和土地利用及人类活动等的影响下其状态会由某一稳态转变到另一稳态。由于环境压力的复杂性、非线性、随机性等特征，往往导致状态转变表现为非线性、突变、跃变等特点。准确界定系统状态跃变的拐点或阈值点存在很大的挑战，而捕捉接近临界拐点前的生态系统结构和属性上的变化特征作为早期预警信号是切实可行的。早期预警信号理论经历理论框架构建、方法确立、机理认知等近半个多世纪的探索，已经由最初的通过仅依赖检测临界点恢复力的速率减慢、方差增加、系统自相关增强等统计学信号过度到更加多样化的检测方法，如检测系统组分属性的变化特征，诊断系统组分各属性之间的关系变化，系统组分的性状变化、系统组分网络结构变化等等，并且试图整合多信号提高预警的精确性。利用来自自然生态系统的长时间高密度数据集和空间代替时间的数据集，基于多度及性状信号的早期预警，结合稳定性、临界恢复力的减速、以及统计参数的指示作用对系统跃变进行早期诊断和预警是预测生态学的主旨。早期预警信号的深入研究不仅能够完善已有理论的不足，同时还能够为生态系统的保护和管理提供切实有效的理论指导。     

陆地生态系统临界转换理论及其生态学机制研究进展

随着气候变化和人类活动对陆地生态系统双重扰动的不断加剧,越来越多的研究已经意识到生态系统结构和功能会发生难以预知的突变,并且恢复起来需要很长时间.开发判别典型生态系统临界转换的早期预警模型及理解其生态学机制成为生态学研究的热点.目前,基于跨越多个时空尺度的理论和实验研究,提出了多种预警陆地生态系统临界转换的理论框架和指...     

Climate‐mediated changes in marine ecosystem regulation during El Niño

The degree to which ecosystems are regulated through bottom‐up, top‐down, or direct physical processes represents a long‐standing issue in ecology, with important consequences for resource management and conservation. In marine ecosystems, the role of bottom‐up and top‐down forcing has been shown to vary over spatio‐temporal scales, often linked to highly variable and heterogeneously distributed environmental conditions. Ecosystem dynamics in the Northeast Pacific have been suggested to be predominately bottom‐up regulated. However, it remains unknown to what extent top‐down regulation occurs, or whether the relative importance of bottom‐up and top‐down forcing may shift in response to climate change. In this study, we investigate the effects and relative importance of bottom‐up, top‐down, and physical forcing during changing climate conditions on ecosystem regulation in the Southern California Current System (SCCS) using a generalized food web model. This statistical approach is based on nonlinear threshold models and a long‐term data set (~60 years) covering multiple trophic levels from phytoplankton to predatory fish. We found bottom‐up control to be the primary mode of ecosystem regulation. However, our results also demonstrate an alternative mode of regulation represented by interacting bottom‐up and top‐down forcing, analogous to wasp‐waist dynamics, but occurring across multiple trophic levels and only during periods of reduced bottom‐up forcing (i.e., weak upwelling, low nutrient concentrations, and primary production). The shifts in ecosystem regulation are caused by changes in ocean‐atmosphere forcing and triggered by highly variable climate conditions associated with El Niño. Furthermore, we show that biota respond differently to major El Niño events during positive or negative phases of the Pacific Decadal Oscillation (PDO), as well as highlight potential concerns for marine and fisheries management by demonstrating increased sensitivity of pelagic fish to exploitation during El Niño.     

Grassland to shrubland state transitions enhance carbon sequestration in the northern Chihuahuan Desert

The replacement of native C₄‐dominated grassland by C₃‐dominated shrubland is considered an ecological state transition where different ecological communities can exist under similar environmental conditions. These state transitions are occurring globally, and may be exacerbated by climate change. One consequence of the global increase in woody vegetation may be enhanced ecosystem carbon sequestration, although the responses of arid and semiarid ecosystems may be highly variable. During a drier than average period from 2007 to 2011 in the northern Chihuahuan Desert, we found established shrubland to sequester 49 g C m^?2 yr^?1 on average, while nearby native C₄ grassland was a net source of 31 g C m^?2 yr^?1 over this same period. Differences in C exchange between these ecosystems were pronounced – grassland had similar productivity compared to shrubland but experienced higher C efflux via ecosystem respiration, while shrubland was a consistent C sink because of a longer growing season and lower ecosystem respiration. At daily timescales, rates of carbon exchange were more sensitive to soil moisture variation in grassland than shrubland, such that grassland had a net uptake of C when wet but lost C when dry. Thus, even under unfavorable, drier than average climate conditions, the state transition from grassland to shrubland resulted in a substantial increase in terrestrial C sequestration. These results illustrate the inherent tradeoffs in quantifying ecosystem services that result from ecological state transitions, such as shrub encroachment. In this case, the deleterious changes to ecosystem services often linked to grassland to shrubland state transitions may at least be partially offset by increased ecosystem carbon sequestration.     

Perturbation and abrupt shift in trophic control of biodiversity and productivity

Ecology is founded on the view that ecosystem properties like biodiversity and productivity change smoothly with changing environmental conditions. However, emerging theory predicts that environmental change may cause abrupt shifts to alternate states. In many ecosystems, top predators play a pivotal role in controlling plant productivity and diversity. Yet it remains uncertain if altering this control shifts systems to alternate states. I report on a test of the hypothesis that loss of predator control of ecosystem function causes abrupt state changes in diversity and productivity. In this meadow ecosystem, predators enhance plant diversity by causing a highly productive, competitively dominant plant species to be suppressed by herbivores. Experimental predator removal caused rapid proliferation of the competitively dominant plant. Moreover, temporally staggered predator reintroductions failed to restore the ecosystem. This loss of resilience confirmed that the ecosystem crossed a critical threshold and entrained into an alternate state.     

Evidence of alternate attractors from a whole-ecosystem regime shift experiment

Ecosystems sometimes shift between different states or dynamic regimes. Theory attributes these shifts to multiple ecosystem attractors. However, documenting multiple ecosystem attractors is difficult, particularly at spatial and temporal scales relevant to ecosystem management. We manipulated the fish community of a lake with the goal of causing trophic cascades and shifting the food web from a planktivore-dominated state to an alternate piscivore-dominated state. We evaluated evidence that the shifts in the fish community comprise alternate attractors using two complementary approaches. First, we calculated phase space trajectories to visualize the shift between attractors. Second, we computed generalized autoregressive conditional heteroskedasticity (GARCH) models and the Brock–Dechert–Scheinkman (BDS) test for linearity. The reconstructed phase space trajectories show the system departing a point attractor, entering a limit cycle, and then shifting to a new point attractor. The GARCH and BDS results indicate that linear explanations are not sufficient to explain the observed patterns. The results provide evidence for alternate attractors based on high-frequency time series of field measurements.     

Non-consumptive effects of predatory mites on thrips and its host plant

Recent reviews on trait-mediated interactions in food webs suggest that trait-mediated effects are as important in triggering top–down trophic cascades as are density-mediated effects. Trait-mediated interactions between predator and prey result from non-consumptive predator effects changing behavioural and/or life history traits of prey. However, in biological control the occurrence of trait-mediated interactions between predators, prey and plants has been largely ignored. Here, we show that non-consumptive predator effects on prey cascade down to the plant in an agro-ecological food chain. The study system consisted of the predatory mites P. persimilis and N. californicus , the herbivorous non-target prey western flower thrips F. occidentalis and the host plant bean. Irrespective of predator species and risk posed to prey, the presence of predator eggs led to increased ambulation, increased mortality and decreased oviposition of thrips. Furthermore, the presence of predator eggs reduced leaf damage caused by thrips. To our knowledge this is the first experimental evidence suggesting a positive trophic cascade triggered by non-consumptive predator effects on non-target prey in an augmentative biological control system.     

A Modeling Study of T-Type Ca Channel Gating and Modulation by L-Cysteine in Rat Nociceptors

L-cysteine (L-cys) increases the amplitude of T-type Ca²⁺ currents in rat T-rich nociceptor-like dorsal root ganglia neurons. The modulation of T-type Ca²⁺ channel gating by L-cys was studied by fitting Markov state models to whole-cell currents recorded from T-rich neurons. The best fitting model tested included three resting states and inactivation from the second resting state and the open state. Inactivation and the final opening step were voltage-independent, whereas transitions between the resting states and deactivation were voltage-dependent. The transition rates between the first two resting states were an order of magnitude faster than those between the second and third resting states, and the voltage-dependency of forward transitions through resting states was two to three times greater than for analogous backward transitions. Analysis with the best fitting model suggested that L-cys increases current amplitude mainly by increasing the transition rate from resting to open and decreasing the transition rate from open to inactivated. An additional model was developed that could account for the bi-exponential time course of recovery from inactivation of the currents and the high frequency of blank sweeps in single channel recordings. This model detected basically the same effects of L-cys on channel gating as the best fitting model.     

On the influence of food quality in consumer–resource interactions

While nutrients are an important regulating factor in food webs, no theoretical studies have examined limits to consumer growth imposed by nutrient concentrations (i.e. food quality) of their prey. Empirical studies have suggested that nutrients may play a role in limiting assimilation efficiencies of herbivores. Using a simple food chain model, I find that prey nutrient concentration does directly influence the growth rate of consumers and potentially increase the stability of consumer–resource interactions. This suggests that the strength of trophic cascades and the relative importance of top–down versus bottom–up control in food webs is significantly influenced by nutrient availability in food resources of consumers. Additionally, the results imply that increases in resource input may cause a change in which resource is limiting and thereby negate any potential "paradox of enrichment".     

Combined Fishing and Climate Forcing in the Southern Benguela Upwelling Ecosystem: An End-to-End Modelling Approach Reveals Dampened Effects

The effects of climate and fishing on marine ecosystems have usually been studied separately, but their interactions make ecosystem dynamics difficult to understand and predict. Of particular interest to management, the potential synergism or antagonism between fishing pressure and climate forcing is analysed in this paper, using an end-to-end ecosystem model of the southern Benguela ecosystem, built from coupling hydrodynamic, biogeochemical and multispecies fish models (ROMS-N₂P₂Z₂D₂-OSMOSE). Scenarios of different intensities of upwelling-favourable wind stress combined with scenarios of fishing top-predator fish were tested. Analyses of isolated drivers show that the bottom-up effect of the climate forcing propagates up the food chain whereas the top-down effect of fishing cascades down to zooplankton in unfavourable environmental conditions but dampens before it reaches phytoplankton. When considering both climate and fishing drivers together, it appears that top-down control dominates the link between top-predator fish and forage fish, whereas interactions between the lower trophic levels are dominated by bottom-up control. The forage fish functional group appears to be a central component of this ecosystem, being the meeting point of two opposite trophic controls. The set of combined scenarios shows that fishing pressure and upwelling-favourable wind stress have mostly dampened effects on fish populations, compared to predictions from the separate effects of the stressors. Dampened effects result in biomass accumulation at the top predator fish level but a depletion of biomass at the forage fish level. This should draw our attention to the evolution of this functional group, which appears as both structurally important in the trophic functioning of the ecosystem, and very sensitive to climate and fishing pressures. In particular, diagnoses considering fishing pressure only might be more optimistic than those that consider combined effects of fishing and environmental variability.     

Ecological effects of longline fishing and climate change on the pelagic ecosystem off eastern Australia

Pelagic longline fisheries target (or catch incidently) large apex predators in the open ocean (e.g. tunas, billfish and sharks) and have the potential to disrupt the ecosystem functionality if these predators exert strong top–down control. In contrast, warming of oceans from climate change may increase bottom–up effects from increases in primary productivity. An ecosystem model of a large pelagic ecosystem off eastern Australia was constructed to explore the potential ecological effects of climate change and longlining by Australia’s Eastern Tuna and Billfish Fishery. The model reproduced historic biomass and fishery catch trends from 1952 to 2006 for seven functional groups. Simulated changes in fishing effort and fishing mortality rate on individual target species from 2008 to 2018 resulted in only modest (<20%) changes in the biomass of target species and their direct predators or competitors. A simulated increase in phytoplankton biomass due to climate change resulted in only small increases (<11%) in the biomass of all groups. However, climate-related changes to the biomass of micronekton fish (−20%) and cephalopods (+50%) resulted in trophic cascades. Our results suggest there may be ecological redundancy among high trophic level predators since they share a diverse suite of prey and collectively only represent <1% of the total system biomass. In contrast, micronekton fishes and cephalopods have high biomasses and high production and consumption rates and are important as both prey and predators. They appear to exert ‘wasp–waist’ control of the ecosystem rather than top–down or bottom–up processes reported to drive other pelagic systems.     

Trophic cascades in the bryosphere: the impact of global change factors on top‐down control of cyanobacterial N2‐fixation

Trophic cascades in which predators regulate densities of organisms at lower trophic levels are important drivers of population dynamics, but effects of trophic cascades on ecosystem‐level fluxes and processes, and the conditions under which top‐down control is important, remain unresolved. We manipulated the structure of a food web in boreal feather mosses and found that moss‐inhabiting microfauna exerted top‐down control of N₂‐fixation by moss‐associated cyanobacteria. However, the presence of higher trophic levels alleviated this top‐down control, likely through feeding on bacterivorous microfauna. These effects of food‐web structure on cyanobacterial N₂‐fixation were dependent on global change factors and strongly suppressed under N fertilisation. Our findings illustrate how food web interactions and trophic cascades can regulate N cycling in boreal ecosystems, where carbon uptake is generally strongly N‐limited, and shifting trophic control of N cycling under global change is therefore likely to impact ecosystem functioning.     

Effects of environmental warming and drought on size‐structured soil food webs

Soil systems maintain important ecosystem processes crucial for plant life and food production. Especially agricultural systems are strongly affected by climate change due to low vegetation cover associated with high temperatures and drought. Nevertheless, the response of soil systems to climate change is little explored. We used microcosms with a simplified soil community to address effects of climate change using independent temperature and dryness gradients and addressed their effects on top–down control and litter decomposition. The community consisted of maize litter as a basal resource, fungi, springtails and as top predators mites and centipedes. As the body‐size structure is of high importance for communities, we included differently‐sized springtails and predator species. After seven weeks, the experiment was terminated, and the impact of climate change on direct feeding interactions and indirect effects across trophic levels was analysed. With increasing temperature and dryness, consumption rates increased, thereby amplifying the negative influence of consumer populations on their resources. Hence, these climate‐change variables increased the top–down control of 1) predators (mainly mites) on springtails and 2) fungi on litter decomposition. In addition, we found that the climate‐change variables strengthened trophic cascades from predators on fungi whose density was thus increasingly decoupled from top–down control by their springtail consumers. Their increased decomposition rates are of high importance for carbon cycling and may result in accelerated nutrient turnover. In conclusion, our results suggest that climate change may strongly influence the structure and functioning of soil systems by strengthening consumption rates and trophic cascades, which will have far reaching consequences for the nutrient turnover and productivity of agricultural ecosystems.     

Trophic amplification of climate warming

Ecosystems can alternate suddenly between contrasting persistent states due to internal processes or external drivers. It is important to understand the mechanisms by which these shifts occur, especially in exploited ecosystems. There have been several abrupt marine ecosystem shifts attributed either to fishing, recent climate change or a combination of these two drivers. We show that temperature has been an important driver of the trophodynamics of the North Sea, a heavily fished marine ecosystem, for nearly 50 years and that a recent pronounced change in temperature established a new ecosystem dynamic regime through a series of internal mechanisms. Using an end-to-end ecosystem approach that included primary producers, primary, secondary and tertiary consumers, and detritivores, we found that temperature modified the relationships among species through nonlinearities in the ecosystem involving ecological thresholds and trophic amplifications. Trophic amplification provides an alternative mechanism to positive feedback to drive an ecosystem towards a new dynamic regime, which in this case favours jellyfish in the plankton and decapods and detritivores in the benthos. Although overfishing is often held responsible for marine ecosystem degeneration, temperature can clearly bring about similar effects. Our results are relevant to ecosystem-based fisheries management (EBFM), seen as the way forward to manage exploited marine ecosystems.     

Review on detection of critical transition in ecosystems

当一个存在多稳态的生态系统临近突变阈值点时, 外界条件即使发生一个微小变化, 也会引发生态系统的剧烈响应, 使之进入结构和功能截然不同的另一稳定状态, 这种现象称为重大突变(critical transition)。重大突变所导致的稳态转换总是伴随着生态系统服务的急剧变化, 可能对人类可持续发展产生重大影响。预测生态系统突变的发生非常困难, 但科学家在此领域的大量研究结果表明, 通过监测一些通用指标可以判断生态系统是否不断临近重大突变阈值点, 进而可以进行生态系统重大突变预警。该文对近年来生态系统重大突变检测领域所取得的成果进行总结与归纳, 论述了生态系统重大突变的产生机制及其后果, 介绍了生态系统突变预警信号提取的理论基础, 从时间和空间两个维度总结了近年来生态系统重大突变预警信号的提取方法, 概述了当前研究面临的挑战, 指出生态系统突变预警信号的检测应充分利用时空动态数据, 并且联合多个指标, 从多个角度进行综合预警, 此外, 还应重视生态系统结构与重大突变之间的关系, 增强生态系统突变预警能力。     

Moderate disturbances accelerate forest transition dynamics under climate change in the temperate–boreal ecotone of eastern North America

Several temperate tree species are expected to migrate northward and colonize boreal forests in response to climate change. Tree migrations could lead to transitions in forest types, but these could be influenced by several non‐climatic factors, such as disturbances and soil conditions. We analysed over 10,000 forest inventory plots, sampled from 1970 to 2018 in meridional Québec, Canada, to identify what environmental conditions promote or prevent regional‐scale forest transitions. We used a continuous‐time multi‐state Markov model to quantify the probabilities of transitions between forest states (temperate, boreal, mixed, pioneer) as a function of climate (mean temperature and climate moisture index during the growing season), soil conditions (pH and drainage) and disturbances (severity levels of natural disturbances and logging). We further investigate how different disturbance types and severities impact forests' short‐term transient dynamics and long‐term equilibrium using properties of Markov transition matrices. The most common transitions observed during the study period were from mixed to temperate states, as well as from pioneer to boreal forests. In our study, transitions were mainly driven by natural and anthropogenic disturbances and secondarily by climate, whereas soil characteristics exerted relatively minor constraints. While major disturbances only promoted transitions to the pioneer state, moderate disturbances increased the probability of transition from mixed to temperate states. Long‐term projections of our model under the current environmental conditions indicate that moderate disturbances would promote a northward shift of the temperate forest. Moreover, disturbances reduced turnover and convergence time for all transitions, thereby accelerating forest dynamics. Contrary to our expectation, mixed to temperate transitions were not driven by temperate tree recruitment but by mortality and growth. Overall, our results suggest that moderate disturbances could catalyse rapid forest transitions and accelerate broad‐scale biome shifts.     

Precursors of state transitions in stochastic systems with delay

Ecosystem dynamics may exhibit alternative stable states induced by positive feedbacks between the state of the system and environmental drivers. Bistable systems are prone to abrupt shifts from one state to another in response to even small and gradual changes in external drivers. These transitions are often catastrophic and difficult to predict by analyzing the mean state of the system. Indicators of the imminent occurrence of phase transitions can serve as important tools to warn ecosystem managers about an imminent transition before the bifurcation point is actually reached. Thus, leading indicators of phase transitions can be used either to prepare for or to prevent the occurrence of a shift to the other state. In recent years, theories of leading indicators of ecosystem shift have been developed and applied to a variety of ecological models and geophysical time series. It is unclear, however, how some of these indicators would perform in the case of systems with a delay. Here, we develop a theoretical framework for the investigation of precursors of state shift in the presence of drivers acting with a delay. We discuss how the effectiveness of leading indicators of state shift based on rising variance may be affected by the presence of delays. We apply this framework to an ecological model of desertification in arid grasslands.     

Mismatch in microbial food webs: predators but not prey perform better in their local biotic and abiotic conditions

Understanding how trophic levels respond to changes in abiotic and biotic conditions is key for predicting how food webs will react to environmental perturbations. Different trophic levels may respond disproportionately to change, with lower levels more likely to react faster, as they typically consist of smaller‐bodied species with higher reproductive rates. This response could cause a mismatch between trophic levels, in which predators and prey will respond differently to changing abiotic or biotic conditions. This mismatch between trophic levels could result in altered top‐down and bottom‐up control and changes in interaction strength. To determine the possibility of a mismatch, we conducted a reciprocal‐transplant experiment involving Sarracenia purpurea food webs consisting of bacterial communities as prey and a subset of six morphologically similar protozoans as predators. We used a factorial design with four temperatures, four bacteria and protozoan biogeographic origins, replicated four times. This design allowed us to determine how predator and prey dynamics were altered by abiotic (temperature) conditions and biotic (predators paired with prey from either their local or non‐local biogeographic origin) conditions. We found that prey reached higher densities in warmer temperature regardless of their temperature of origin. Conversely, predators achieved higher densities in the temperature condition and with the prey from their origin. These results confirm that predators perform better in abiotic and biotic conditions of their origin while their prey do not. This mismatch between trophic levels may be especially significant under climate change, potentially disrupting ecosystem functioning by disproportionately affecting top‐down and bottom‐up control.