Long-term Perspectives on Lagged Ecosystem Responses to Climate Change: Permafrost in Boreal Peatlands and the Grassland/Woodland Boundary

Changes in climate could have far-reaching consequences for ecosystems sensitive to changes in temperature and precipitation, such as boreal permafrost peatlands and grassland/woodland boundaries. The long-term data from our studies in these ecosystems suggest that transient responses of permafrost and vegetation to climate change may be difficult to predict due to lags and positive feedbacks related to vegetation and disturbance. Boreal permafrost peatlands comprise an ecosystem with strong local controls on microclimate that influence the formation and thaw of permafrost. These local controls may preserve permafrost during the transient stages of climate warming, producing lagged responses. The prairie–forest border region of the northern Great Plains has experienced frequent change and has complex dynamics involving transitions in the grassland composition of prairie and in the degree of woodiness in bordering forests. Fire frequency interacts with fuel loading and tree recruitment in ways that affect the timing and direction of change. Lags and thresholds could lead to sudden large responses to future climate change that are not readily apparent from current vegetation. The creation of adequate models to characterize transient ecosystem changes will require an understanding of the linkages among processes operating at the scale of 10s of meters and over long time periods. Received 14 December 1999; accepted 7 July 2000.     

Tropical terrestrial invertebrates—Where to from here?

There are over one million described invertebrate species on Earth, the majority of which are likely to inhabit the highly biodiverse rain forests around the equator. These are some of the most vulnerable ecosystems on Earth due to the pressures of deforestation and climate change with many of their inhabitants at risk of extinction. Invertebrates play a major role in ecosystem functioning from decomposition and nutrient cycling to herbivory and pollination; however, while our understanding of these roles is improving, we are far from being able to predict the consequences of further deforestation, climate change, and biodiversity loss due to the lack of comparative data and the high proportion of species which remain to be discovered. As we move into an era of increased pressure on old-growth habitats and biodiversity, it is imperative that we understand how changes to invertebrate communities, and the extinction of species, affect ecosystems. Innovative and comprehensive methods that approach these issues are needed. Here, we highlight priorities for future tropical terrestrial invertebrate research such as the efficiency of sustainable land management, exploration of innovative methods for better understanding of invertebrate ecology and behavior, and quantifying the role of invertebrates in ecosystem functioning.     

Environmental paleomicrobiology: using DNA preserved in aquatic sediments to its full potential

In-depth knowledge about spatial and temporal variation in microbial diversity and function is needed for a better understanding of ecological and evolutionary responses to global change. In particular, the study of microbial ancient DNA preserved in sediment archives from lakes and oceans can help us to evaluate the responses of aquatic microbes in the past and make predictions about future biodiversity change in those ecosystems. Recent advances in molecular genetic methods applied to the analysis of historically deposited DNA in sediments have not only allowed the taxonomic identification of past aquatic microbial communities but also enabled tracing their evolution and adaptation to episodic disturbances and gradual environmental change. Nevertheless, some challenges remain for scientists to take full advantage of the rapidly developing field of paleo-genetics, including the limited ability to detect rare taxa and reconstruct complete genomes for evolutionary studies. Here, we provide a brief review of some of the recent advances in the field of environmental paleomicrobiology and discuss remaining challenges related to the application of molecular genetic methods to study microbial diversity, ecology, and evolution in sediment archives. We anticipate that, in the near future, environmental paleomicrobiology will shed new light on the processes of microbial genome evolution and microbial ecosystem responses to quaternary environmental changes at an unprecedented level of detail. This information can, for example, aid geological reconstructions of biogeochemical cycles and predict ecosystem responses to environmental perturbations, including in the context of human-induced global changes.     

Pathways to bridge the biophysical realism gap in ecosystem services mapping approaches

The mapping of ecosystem service supply has become quite common in ecosystem service assessment practice for terrestrial ecosystems, but land cover remains the most common indicator for ecosystems ability to deliver ecosystem services. For marine ecosystems, practice is even less advanced, with a clear deficit in spatially-explicit assessments of ecosystem service supply. This situation, which generates considerable uncertainty in the assessment of ecosystems’ ability to support current and future human well-being, contrasts with increasing understanding of the role of terrestrial and marine biodiversity for ecosystem functioning and thereby for ecosystem services. This paper provides a synthesis of available approaches, models and tools, and data sources, that are able to better link ecosystem service mapping to current understanding of the role of ecosystem service providing organisms and land/seascape structure in ecosystem functioning. Based on a review of literature, models and associated geo-referenced metrics are classified according to the way in which land or marine use, ecological processes and especially biodiversity effects are represented. We distinguish five types of models: proxy-based, phenomenological, niche-based, trait-based and full-process. Examples from each model type are presented and data requirements considered. Our synthesis demonstrates that the current understanding of the role of biota in ecosystem services can effectively be incorporated into mapping approaches and opens avenues for further model development using hybrid approaches tailored to available resources. We end by discussing ways to resolve sources of uncertainty associated with model representation of biotic processes and with data availability.     

Predicting Novel Riparian Ecosystems in a Changing Climate

Rapid changes in global climate are likely to alter species assemblages and environmental characteristics resulting in novel ecosystems. The ability to predict characteristics of future ecosystems is crucial for environmental planning and the development of effective climate change adaptation strategies. This paper presents an approach for envisioning novel ecosystems in future climates. Focusing on riparian ecosystems, we use qualitative process models to predict likely abiotic and biotic changes in four case study systems: tropical coastal floodplains, temperate streams, high mountain streams and urban riparian zones. We concentrate on functional groups rather than individual species and consider dispersal constraints and the capacity for genetic adaptation. Our scenarios suggest that climatic changes will reduce indigenous diversity, facilitate non-indigenous invasion (especially C4 graminoids), increase fragmentation and result in simplified and less distinctive riparian ecosystems. Compared to models based on biota-environment correlations, process models built on mechanistic understanding (like Bayesian belief networks) are more likely to remain valid under novel climatic conditions. We posit that predictions based on species’ functional traits will facilitate regional comparisons and can highlight effects of climate change on ecosystem structure and function. Ecosystems that have experienced similar modification to that expected under climate change (for example, altered flow regimes of regulated rivers) can be used to help inform and evaluate predictions. By manipulating attributes of these system models (for example, magnitude of climatic changes or adaptation strategies used), implications of various scenarios can be assessed and optimal management strategies identified.     

全球环境变化对典型生态系统的影响研究:现状、挑战与发展趋势

随着全球环境变化和人类活动对生态系统影响的日益加深,生态系统结构和功能发生强烈变化,生态系统提供各类资源和服务的能力在显著下降。在这种背景下,全面认识生态系统的结构功能与全球环境变化的关系已成为当前生态学研究的热点之一。本文综述了全球环境变化对典型生态系统(包括森林生态系统、河口湿地生态系统、城市生态系统)影响以及全球环境变化适应的研究现状,分析研究面临的困难及挑战。在此基础上,提出对未来研究发展趋势的展望。在森林生态系统与全球环境变化研究上,未来应重视能更好模拟现实情景的、多因子、长期的全球环境变化控制试验,并注重不同生物地球化学循环之间的耦合作用。在湿地生态系统与全球环境变化研究上,未来应加强氮沉降、硫沉降及盐水入侵对湿地生态系统碳氮循环的影响,明晰滨海湿地的蓝碳功能,加强极端气候和人类干扰影响下湿地生态系统结构和功能变化及恢复力的研究。在城市生态系统与全球环境变化研究上,未来应深化城市生物地球化学循环机制研究,实现城市生态系统的人本需求侧重与转向,并开展典型地区长期、多要素综合响应研究。在全球环境变化适应研究上,未来应构架定量化、跨尺度的适应性评价体系,加强典型区域/部门的适应性研究以及适应策略实施的可行性研究,注重适应与减缓对策的关联研究及实施的风险评估。期望本综述为我国生态系统与全球环境变化研究提供一些参考。     

Analyzing trophic ecosystem functions with the interaction functional space

Quantifying the vulnerability of ecosystems to global change requires a better understanding of how trophic ecosystem functions emerge. So far, trophic ecosystem functions have been studied from the perspective of either functional diversity or network ecology. To integrate these two perspectives, we propose the interaction functional space (IFS) a conceptual framework to simultaneously analyze the effects of traits and interactions on trophic functions. We exemplify the added value of our framework for seed dispersal and wood decomposition and show how species interactions influence the relationship between functional trait diversity and trophic functions. We propose future applications for a range of functions where the IFS can help to elucidate mechanisms underpinning trophic functions and facilitate understanding of functional changes in ecosystems amidst global change.     

From Transient to Steady-state Response of Ecosystems to Atmospheric CO2-enrichment and Global Climate Change: Conceptual Challenges and Need for an Integrated Approach

Evidence continues to accumulate that humans are significantly increasing atmospheric CO₂ concentrations, resulting in unprecedented changes in the global climate system. Experimental manipulations of terrestrial ecosystems and their components have greatly increased our understanding of short-term responses to these global perturbations and have provided valuable input to ecosystem, dynamic vegetation, and global scale models. However, concerns exist that these initial experimental responses may be transitory, thereby limiting our ability to extrapolate short-term experimental responses to infer longer-term effects. To do these extrapolations, it will be necessary to understand changes in response patterns over time, including alterations in the magnitude, direction, and rate of change of the responses. These issues represent one of our largest challenges in accurately predicting longer-term changes in ecosystems and associated feedbacks to the climate system. Key issues that need to be considered when designing future experiments or refining models include: linear vs. non-linear responses, direct vs. indirect effects, lags in response, acclimation, resource limitation, homeostasis, buffers, thresholds, ecosystem stoichiometry, turnover rates and times, and alterations in species composition. Although experimental and landscape evidence for these response patterns exist, extrapolating longer-term response patterns from short-term experiments will ultimately require a unified multidisciplinary approach, including better communication and collaboration between theoreticists, experimentalists and modelers.     

Historical climate‐human‐ecosystem interaction in East Africa: a review

The palaeoecological visibility of historical human impact on natural ecosystems in tropical East Africa is strongly impeded by an overriding dominant signature of climate change at decadal‐to‐millennial time scales. Better knowledge of the relative magnitude and timing of present and past human impact and climate variability is, however, instrumental to properly assess the resilience, and recovery potential, of East Africa's natural ecosystems. Here, we briefly review comprehensive previous attempts to assess past ecosystem responses to climate change and human impact. We further discuss some key issues of climate‐human‐ecosystem relationships in a multidisciplinary framework and address some future challenges and outcomes, which may pave the way to a better understanding of past climate‐human‐ecosystem interaction‐ in tropical Africa.     

Predicting the likely response of data‐poor ecosystems to climate change using space‐for‐time substitution across domains

Predicting ecological response to climate change is often limited by a lack of relevant local data from which directly applicable mechanistic models can be developed. This limits predictions to qualitative assessments or simplistic rules of thumb in data‐poor regions, making management of the relevant systems difficult. We demonstrate a method for developing quantitative predictions of ecological response in data‐poor ecosystems based on a space‐for‐time substitution, using distant, well‐studied systems across an inherent climatic gradient to predict ecological response. Changes in biophysical data across the spatial gradient are used to generate quantitative hypotheses of temporal ecological responses that are then tested in a target region. Transferability of predictions among distant locations, the novel outcome of this method, is demonstrated via simple quantitative relationships that identify direct and indirect impacts of climate change on physical, chemical and ecological variables using commonly available data sources. Based on a limited subset of data, these relationships were demonstrably plausible in similar yet distant (>2000 km) ecosystems. Quantitative forecasts of ecological change based on climate‐ecosystem relationships from distant regions provides a basis for research planning and informed management decisions, especially in the many ecosystems for which there are few data. This application of gradient studies across domains – to investigate ecological response to climate change – allows for the quantification of effects on potentially numerous, interacting and complex ecosystem components and how they may vary, especially over long time periods (e.g. decades). These quantitative and integrated long‐term predictions will be of significant value to natural resource practitioners attempting to manage data‐poor ecosystems to prevent or limit the loss of ecological value. The method is likely to be applicable to many ecosystem types, providing a robust scientific basis for estimating likely impacts of future climate change in ecosystems where no such method currently exists.     

Eight questions about invasions and ecosystem functioning

I pose eight questions central to understanding how biological invasions affect ecosystems, assess progress towards answering those questions and suggest ways in which progress might be made. The questions concern the frequency with which invasions affect ecosystems; the circumstances under which ecosystem change is most likely; the functions that are most often affected by invaders; the relationships between changes to ecosystems, communities, and populations; the long-term responses of ecosystems to invasions; interactions between biological invasions and other anthropogenic activities and the difficulty of managing undesirable impacts of non-native species. Some questions have been answered satisfactorily, others require more data and thought, and others might benefit from being reformulated or abandoned. Actions that might speed progress include careful development of trait-based approaches; strategic collection and publication of new data, including more frequent publication of negative results; replacement of expert opinion with hard data where needed; careful consideration of whether questions really need to be answered, especially in cases where answers are being provided for managers and policy-makers; explicit attention to and testing of the domains of theories; integrating invasions better into an ecosystem context; and remembering that our predictive ability is limited and will remain so for the foreseeable future.     

Exploring the dynamics of ecological indicators using food web models fitted to time series of abundance and catch data

Previously, standardized snap-shot models of the Southern Benguela (1980–1989), Southern Humboldt (1992) and Southern Catalan Sea (1994) ecosystems were examined and found to facilitate assessment of ecosystem characteristics related to the gradient in exploitation status of the ecosystems; highest level of exploitation in the South Catalan Sea (North-western Mediterranean), high in the Southern Humboldt and lower in the Southern Benguela. Subsequently, these models were calibrated and fitted using available catch, fishing effort/mortality and abundance data series and incorporated environmental and internal drivers. This study furthers the previous comparative analyses by comparing changes in ecosystem structure using a selection of ecosystem indicators from the calibrated models and assessing how these indicators change over time in these three contrasting ecosystems. Indicators examined include community turnover rates (production/biomass), trophic level of landings and the community, biodiversity indicators, ratios of predatory/forage fish and pelagic/demersal fish biomass, catch ratios, and network analysis indicators. Using the set of model-derived indicators, the three ecosystems were ranked in terms of exploitation level. This ranking was performed using the values of these indicators in recent years (ecosystem state) as well as their trends over time (ecosystem trend). The non-parametric Kruskal–Wallis and Median tests were used to test for significance of the difference between indicators from the three ecosystems in the last 5 years of the simulation to compare present ecosystem states. We compared the slope of the lineal trend and its significance between ecosystems using the generalized least-squares regression taking auto-correlation into consideration to analyse ecosystem trends. The indicators that capture better the high impacts of fishing prevalent in the Mediterranean and Humboldt ecosystems, and the more conservative exploitation of the Southern Benguela, are the fish/invertebrates biomass and catch ratio, the demersal/pelagic fish biomass and catch ratio (depending on the ecosystem and the fishery being developed), flows to detritus, and the mean trophic level of the community (when large, poorly quantified groups such as zooplankton and detritus are excluded). This study suggests that the best option for classifying ecosystems according to the impact of fishing is to consider a broad range of indicators to understand how and why an ecosystem is responding to particular environmental or fishing drivers (or more likely a combination of these). Our results highlight the importance of including indicators capturing trends over time as well as recent ecosystem states. We also identified 23 pairs of indicators that correlated similarly in the three ecosystems (they showed a significant correlation with same sign). Further comparisons may contribute towards generalization of this list, progressing towards a better understanding of the behaviour of ecological indicators.     

The human dimension of fire regimes on Earth

Humans and their ancestors are unique in being a fire-making species, but 'natural' (i.e. independent of humans) fires have an ancient, geological history on Earth. Natural fires have influenced biological evolution and global biogeochemical cycles, making fire integral to the functioning of some biomes. Globally, debate rages about the impact on ecosystems of prehistoric human-set fires, with views ranging from catastrophic to negligible. Understanding of the diversity of human fire regimes on Earth in the past, present and future remains rudimentary. It remains uncertain how humans have caused a departure from 'natural' background levels that vary with climate change. Available evidence shows that modern humans can increase or decrease background levels of natural fire activity by clearing forests, promoting grazing, dispersing plants, altering ignition patterns and actively suppressing fires, thereby causing substantial ecosystem changes and loss of biodiversity. Some of these contemporary fire regimes cause substantial economic disruptions owing to the destruction of infrastructure, degradation of ecosystem services, loss of life, and smoke-related health effects. These episodic disasters help frame negative public attitudes towards landscape fires, despite the need for burning to sustain some ecosystems. Greenhouse gas-induced warming and changes in the hydrological cycle may increase the occurrence of large, severe fires, with potentially significant feedbacks to the Earth system. Improved understanding of human fire regimes demands: (1) better data on past and current human influences on fire regimes to enable global comparative analyses, (2) a greater understanding of different cultural traditions of landscape burning and their positive and negative social, economic and ecological effects, and (3) more realistic representations of anthropogenic fire in global vegetation and climate change models. We provide an historical framework to promote understanding of the development and diversification of fire regimes, covering the pre-human period, human domestication of fire, and the subsequent transition from subsistence agriculture to industrial economies. All of these phases still occur on Earth, providing opportunities for comparative research.     

陆地生态系统植被氮磷化学计量研究进展

因化学功能的耦合和元素的不可替代性, 植物对N、P的需求和利用存在严格的比例。植物N、P化学计量在不同功能群、生长地区、生长季、器官之间以及环境梯度下存在明显的变化规律。多数研究从N、P浓度、N:P及N、P间异速指数等角度分析了植物化学计量变化规律, 并探讨其在全球范围内的具体数值。为增进对植物响应全球变化的理解, 该文综述了N、P化学计量的影响因素及其机理的最新研究进展, 并指出未来拟重点研究的方向。     

Modelling climate‐change‐induced nonlinear thresholds in cephalopod population dynamics

A significant global challenge lies in our current inability to anticipate, and therefore prepare for, critical ecological thresholds (i.e. tipping points in ecosystems). This deficit stems largely from an inadequate understanding of the many complex interactions between species and the environment at the ecosystem level, and the paucity of mechanistic models relating environment to population dynamics at the species level. In marine ecosystems, abundant, short‐lived and fast‐growing species such as anchovies or squids, consistently function as ‘keystone’ groups whose population dynamics affect entire ecosystems. Increasing exploitation coupled with climate change impacts has the potential to affect these ecological groups and consequently, the entire marine ecosystem. There are currently very few models that predict the impact of climate change on these keystone groups. Here we use a combination of individual‐based bioenergetics and stage‐structured population models to characterize the fundamental capacity of cephalopods to respond to climate change. We demonstrate the potential for, and mechanisms behind, two unfavourable climate‐change‐induced thresholds in future population dynamics. Although one threshold was the direct consequence of a decrease in incubation time caused by ocean warming, the other threshold was linked to survivorship, implying the possibility of management through a modification of fishing mortality. Additional substantive changes in phenology were also predicted, with a possible loss in population resilience. Our results demonstrate the feasibility of predicting complex nonlinear dynamics with a reasonably simplistic mechanistic model, and highlight the necessity of developing such approaches for other species if attempts to moderate the impact of climate change on natural resources are to be effective.     

Predicting Plant Diversity Response to Disturbance: Applicability of the Intermediate Disturbance Hypothesis and Mass Ratio Hypothesis

Predicting the relationships between disturbance, biodiversity and productivity of ecosystems continue to preoccupy ecologists and resource managers. Two hypotheses underpin many of the discussions. The Intermediate Disturbance Hypothesis (IDH), which proposes that biodiversity peaks at intermediate levels of disturbance, is often extended to predict that productivity follows the same response pattern. The Mass Ratio Hypothesis (MRH) proposes that the biological traits of the dominant species are the critical drivers of ecosystem function (e.g., productivity) and that these species increase in biomass rapidly after disturbance then stabilize. As a consequence, species diversity first peaks then declines after disturbance as a few species dominate the site. Both provide a conceptual link among disturbance, species diversity and productivity (an index of ecosystem function). We assessed the current state of empirical support for these two hypotheses with a literature survey and determined if their conformance is related to ecosystem type or site productivity. Conformance of IDH reported in past reviews (considering all ecosystems) ranged from 16 to 21%. This contrasts with our finding that in terrestrial ecosystems conformance to IDH was 46% (22 of 48 studies), 17% studies reported non-compliance, and 23% reported inconclusive results. Most studies explained their results with respect to IDH or MRH. Only two studies were specifically designed to test the validity of IDH or MRH. We conclude that (i) the IDH is mostly applicable to predict species diversity response to disturbance in upland sites of medium to high productivity and the MRH is applicable to organic sites of low productivity; (ii) there is a critical need for more studies specifically designed to test these hypotheses in natural ecosystems using common protocols; and (iii) enhanced understanding of these models will add value in refining management policies and in the selection of meaningful diversity indicators of sustainability.     

Possible effects of global environmental changes on Antarctic benthos: a synthesis across five major taxa

Because of the unique conditions that exist around the Antarctic continent, Southern Ocean (SO) ecosystems are very susceptible to the growing impact of global climate change and other anthropogenic influences. Consequently, there is an urgent need to understand how SO marine life will cope with expected future changes in the environment. Studies of Antarctic organisms have shown that individual species and higher taxa display different degrees of sensitivity to environmental shifts, making it difficult to predict overall community or ecosystem responses. This emphasizes the need for an improved understanding of the Antarctic benthic ecosystem response to global climate change using a multitaxon approach with consideration of different levels of biological organization. Here, we provide a synthesis of the ability of five important Antarctic benthic taxa (Foraminifera, Nematoda, Amphipoda, Isopoda, and Echinoidea) to cope with changes in the environment (temperature, pH, ice cover, ice scouring, food quantity, and quality) that are linked to climatic changes. Responses from individual to the taxon-specific community level to these drivers will vary with taxon but will include local species extinctions, invasions of warmer-water species, shifts in diversity, dominance, and trophic group composition, all with likely consequences for ecosystem functioning. Limitations in our current knowledge and understanding of climate change effects on the different levels are discussed.     

Key ecological research questions for Central European forests

Forests are under pressure from accelerating global change. To cope with the multiple challenges related to global change but also to further improve forest management we need a better understanding of (1) the linkages between drivers of ecosystem change and the state and management of forest ecosystems as well as their capacity to adapt to ongoing global environmental changes, and (2) the interrelationships within and between the components of forest ecosystems. To address the resulting challenges for the state of forest ecosystems in Central Europe, we suggest 45 questions for future ecological research. We define forest ecology as studies on the abiotic and biotic components of forest ecosystems and their interactions on varying spatial and temporal scales. Our questions cover five thematic fields and correspond to the criteria selected for describing the state of Europe’s forests by policy makers, i.e. biogeochemical cycling, mortality and disturbances, productivity, biodiversity and biotic interactions, and regulation and protection. We conclude that an improved mechanistic understanding of forest ecosystems is essential for the further development of ecosystem-oriented multifunctional forest management in the face of accelerating global change.     

Seasonal,not annual precipitation drives community productivity across ecosystems

Understanding drivers of aboveground net primary production (ANPP) has long been a goal of ecology. Decades of investigation have shown total annual precipitation to be an important determinant of ANPP within and across ecosystems. Recently a few studies at individual sites have shown precipitation during specific seasons of the year can more effectively predict ANPP. Here we determined whether seasonal or total precipitation better predicted ANPP across a range of terrestrial ecosystems, from deserts to forests, using long‐term data from 36 plant communities. We also determined whether ANPP responses were dependent on ecosystem type or plant functional group. We found that seasonal precipitation generally explained ANPP better than total precipitation. Precipitation in multiple parts of the growing season often correlated with ANPP, but rarely interacted with each other. Surprisingly, the amount of variation explained by seasonal precipitation was not correlated with ecosystem type or plant functional group. Overall, examining seasonal precipitation can significantly improve ANPP predictions across a broad range of ecosystems and plant types, with implications for understanding current and future ANPP variation. Further work examining precipitation timing relative to species phenology may further improve our ability to predict ANPP, especially in response to climate change.     

土壤线虫对气候变化的响应研究进展

全球变化对陆地生态系统功能具有重要而深远的影响。陆地生态系统地下部分具有重要的生态功能,其组成及结构对气候变化的响应将进一步减缓或加剧全球化进程。土壤线虫在各类生态系统中分布十分广泛,是地下食物网的重要组分,在维持土壤生物多样性及营养物质循环过程中发挥重要作用,其组成及结构对不同气候变化驱动因子的响应机制与模式不尽相同。增温及降水格局变化主要是通过改变线虫生境而直接影响其种群密度与结构,两者通常表现为正效应且作用效果随处理时间的延长而增强。CO2与大气氮沉降主要是通过影响地上植被,凋落物质量,土壤理化性质等间接过程影响土壤线虫。同时,不同的全球变化因子之间存在着复杂的交互作用,深入理解这些因子之间交互作用对线虫群落的影响模式与机制对于探讨未来气候变化情景下生态统生物多样性及养分循环过程具有重要的理论指导意义。