From observations to experiments in phenology research: investigating climate change impacts on trees and shrubs using dormant twigs

Background and Aims Climate change is advancing the leaf-out times of many plant species and mostly extending the growing season in temperate ecosystems. Laboratory experiments using twig cuttings from woody plant species present an affordable, easily replicated approach to investigate the relative importance of factors such as winter chilling, photoperiod, spring warming and frost tolerance on the leafing-out times of plant communities. This Viewpoint article demonstrates how the results of these experiments deepen our understanding beyond what is possible via analyses of remote sensing and field observation data, and can be used to improve climate change forecasts of shifts in phenology, ecosystem processes and ecological interactions.Scope The twig method involves cutting dormant twigs from trees, shrubs and vines on a single date or at intervals over the course of the winter and early spring, placing them in containers of water in controlled environments, and regularly recording leaf-out, flowering or other phenomena. Prior to or following leaf-out or flowering, twigs may be assigned to treatment groups for experiments involving temperature, photoperiod, frost, humidity and more. Recent studies using these methods have shown that winter chilling requirements and spring warming strongly affect leaf-out and flowering times of temperate trees and shrubs, whereas photoperiod requirements are less important than previously thought for most species. Invasive plant species have weaker winter chilling requirements than native species in temperate ecosystems, and species that leaf-out early in the season have greater frost tolerance than later leafing species.Conclusions This methodology could be extended to investigate additional drivers of leaf-out phenology, leaf senescence in the autumn, and other phenomena, and could be a useful tool for education and outreach. Additional ecosystems, such as boreal, southern hemisphere and sub-tropical forests, could also be investigated using dormant twigs to determine the drivers of leaf-out times and how these ecosystems will be affected by climate change.     

Global shifts towards positive species interactions with increasing environmental stress

The study of positive species interactions is a rapidly evolving field in ecology. Despite decades of research, controversy has emerged as to whether positive and negative interactions predictably shift with increasing environmental stress as hypothesised by the stress‐gradient hypothesis (SGH). Here, we provide a synthesis of 727 tests of the SGH in plant communities across the globe to examine its generality across a variety of ecological factors. Our results show that plant interactions change with stress through an outright shift to facilitation (survival) or a reduction in competition (growth and reproduction). In a limited number of cases, plant interactions do not respond to stress, but they never shift towards competition with stress. These findings are consistent across stress types, plant growth forms, life histories, origins (invasive vs. native), climates, ecosystems and methodologies, though the magnitude of the shifts towards facilitation with stress is dependent on these factors. We suggest that future studies should employ standardised definitions and protocols to test the SGH, take a multi‐factorial approach that considers variables such as plant traits in addition to stress, and apply the SGH to better understand how species and communities will respond to environmental change.     

Quantifying the impact of ecological memory on the dynamics of interacting communities

Ecological memory refers to the influence of past events on the response of an ecosystem to exogenous or endogenous changes. Memory has been widely recognized as a key contributor to the dynamics of ecosystems and other complex systems, yet quantitative community models often ignore memory and its implications.Recent modeling studies have shown how interactions between community members can lead to the emergence of resilience and multistability under environmental perturbations. We demonstrate how memory can be introduced in such models using the framework of fractional calculus. We study how the dynamics of a well-characterized interaction model is affected by gradual increases in ecological memory under varying initial conditions, perturbations, and stochasticity.Our results highlight the implications of memory on several key aspects of community dynamics. In general, memory introduces inertia into the dynamics. This favors species coexistence under perturbation, enhances system resistance to state shifts, mitigates hysteresis, and can affect system resilience both ways depending on the time scale considered. Memory also promotes long transient dynamics, such as long-standing oscillations and delayed regime shifts, and contributes to the emergence and persistence of alternative stable states. Our study highlights the fundamental role of memory in communities, and provides quantitative tools to introduce it in ecological models and analyse its impact under varying conditions.     

Vegetation change from chronic stress events: Detection of the effects of tide gate removal and long‐term drought on a tidal marsh

Question: Chronic stress events are defined as disturbance events that exceed the lifespan of the dominant plant species, fluctuate in intensity and lack abruptness or physical destruction of biomass. Can the effects of chronic stress events be measured on vegetation communities? Did two chronic stress events, the removal of a tide gate and a four year drought, cause a temporary or permanent shift in the vegetation communities of a tidal marsh? Location: Tidal marsh in southeastern United States. Methods: Change in species composition and dominance and community change on a landscape level salinity gradient were measured between time periods ranging from four months to seven years to construct a statistical baseline reference community at freshwater, oligohaline, and mesohaline sections of a tidal marsh. Statistical shifts in the plant community were defined as changes in the plant community that fell outside of the defined baseline reference community. Results: Plant community changes outside of the reference community occurred in 13 out of 378 community comparisons. Removal of the tide gate had a greater effect on interstitial salinity levels than the drought and was most intense in the oligohaline marsh, where between 20 to 45% of the freshwa‐ter/oligohaline community types permanently converted to oligohaline community types. However, community shifts in the freshwater and oligohaline marsh induced by the drought were temporary, lasting from 1 to 3+ years. Neither chronic stress event permanently altered the mesohaline plant communities. Conclusion: The effects of chronic stress events could be detected; an extended historical record of vegetation change (18 years) was necessary to identify community shifts outside of a reference condition of the community and to determine if those shifts were permanent or temporary.     

Means and extremes: building variability into community‐level climate change experiments

Experimental studies assessing climatic effects on ecological communities have typically applied static warming treatments. Although these studies have been informative, they have usually failed to incorporate either current or predicted future, patterns of variability. Future climates are likely to include extreme events which have greater impacts on ecological systems than changes in means alone. Here, we review the studies which have used experiments to assess impacts of temperature on marine, freshwater and terrestrial communities, and classify them into a set of ‘generations’ based on how they incorporate variability. The majority of studies have failed to incorporate extreme events. In terrestrial ecosystems in particular, experimental treatments have reduced temperature variability, when most climate models predict increased variability. Marine studies have tended to not concentrate on changes in variability, likely in part because the thermal mass of oceans will moderate variation. In freshwaters, climate change experiments have a much shorter history than in the other ecosystems, and have tended to take a relatively simple approach. We propose a new ‘generation’ of climate change experiments using down‐scaled climate models which incorporate predicted changes in climatic variability, and describe a process for generating data which can be applied as experimental climate change treatments.     

Impacts of recent climate change on terrestrial flora and fauna: Some emerging Australian examples

The effects of anthropogenic climate change on biodiversity are well known for some high‐profile Australian marine systems, including coral bleaching and kelp forest devastation. Less well‐published are the impacts of climate change being observed in terrestrial ecosystems, although ecological models have predicted substantial changes are likely. Detecting and attributing terrestrial changes to anthropogenic factors is difficult due to the ecological importance of extreme conditions, the noisy nature of short‐term data collected with limited resources, and complexities introduced by biotic interactions. Here, we provide a suite of case studies that have considered possible impacts of anthropogenic climate change on Australian terrestrial systems. Our intention is to provide a diverse collection of stories illustrating how Australian flora and fauna are likely responding to direct and indirect effects of anthropogenic climate change. We aim to raise awareness rather than be comprehensive. We include case studies covering canopy dieback in forests, compositional shifts in vegetation, positive feedbacks between climate, vegetation and disturbance regimes, local extinctions in plants, size changes in birds, phenological shifts in reproduction and shifting biotic interactions that threaten communities and endangered species. Some of these changes are direct and clear cut, others are indirect and less clearly connected to climate change; however, all are important in providing insights into the future state of terrestrial ecosystems. We also highlight some of the management issues relevant to conserving terrestrial communities and ecosystems in the face of anthropogenic climate change.     

Extreme weather events and plant–plant interactions: shifts between competition and facilitation among grassland species in the face of drought and heavy rainfall

Biotic interactions play an important role in ecosystem function and structure in the face of global climate change. We tested how plant–plant interactions, namely competition and facilitation among grassland species, respond to extreme drought and heavy rainfall events. We also examined how the functional composition (grasses, forbs, legumes) of grassland communities influenced the competition intensity for grass species when facing extreme events. We exposed experimental grassland communities of different functional compositions to either an extreme single drought event or to a prolonged heavy rainfall event. Relative neighbour effect, relative crowding and interaction strength were calculated for five widespread European grassland species to quantify competition. Single climatic extremes caused species specific shifts in plant–plant interactions from facilitation to competition or vice versa but the nature of the shifts varied depending on the community composition. Facilitation by neighbouring plants was observed for Arrhenatherum elatius when subjected to drought. Contrarily, the facilitative effect of neighbours on Lotus corniculatus was transformed into competition. Heavy rainfall increased the competitive effect of neighbours on Holcus lanatus and Lotus corniculatus in communities composed of three functional groups. Competitive pressure on Geranium pratense and Plantago lanceolata was not affected by extreme weather events. Neither heavy rainfall nor extreme drought altered the overall productivity of the grassland communities. The complementary responses in competition intensity experienced by grassland species under drought suggest biotic interactions as one stabilizing mechanism for overall community performance. Understanding competitive dynamics under fluctuating resources is important for assessing plant community shifts and degree of stability of ecosystem functions.     

Effects of plant species richness on invasion dynamics, disease outbreaks, insect abundances and diversity

Declining biodiversity represents one of the most dramatic and irreversible aspects of anthropogenic global change, yet the ecological implications of this change are poorly understood. Recent studies have shown that biodiversity loss of basal species, such as autotrophs or plants, affects fundamental ecosystem processes such as nutrient dynamics and autotrophic production. Ecological theory predicts that changes induced by the loss of biodiversity at the base of an ecosystem should impact the entire system. Here we show that experimental reductions in grassland plant richness increase ecosystem vulnerability to invasions by plant species, enhance the spread of plant fungal diseases, and alter the richness and structure of insect communities. These results suggest that the loss of basal species may have profound effects on the integrity and functioning of ecosystems.     

Interactive effects of anthropogenic nitrogen enrichment and climate change on terrestrial and aquatic biodiversity

Biodiversity has been described as the diversity of life on earth within species, among species, and among ecosystems. The rate of biodiversity loss due to human activity in the last 50 years has been more rapid than at any other time in human history, and many of the drivers of biodiversity loss are increasing, including habitat loss, overexploitation, invasive species, climate change, and pollution, including pollution from reactive nitrogen (Nr). Of these stressors, climate change and Nr from anthropogenic activities are causing some of the most rapid changes. Climate change is causing warming trends that result in poleward and elevational range shifts of flora and fauna, and changes in phenology, particularly the earlier onset of spring events and migration, and lengthening of the growing season. Nitrogen (N) enrichment can enhance plant growth, but has been shown to favor, fast-growing, sometimes invasive, species over native species adapted to low N conditions. Although there have been only a few controlled studies on climate change and N interactions, inferences can be drawn from various field observations. For example, in arid ecosystems of southern California, elevated N deposition and changing precipitation patterns have promoted the conversion of native shrub communities to communities dominated by annual non-native grasses. Both empirical studies and modeling indicate that N and climate change can interact to drive losses in biodiversity greater than those caused by either stressor alone. Reducing inputs of anthropogenic Nr may be an effective mitigation strategy for protecting biodiversity in the face of climate change.     

Shifts in plant functional community composition under hydrological stress strongly decelerate litter decomposition

Litter decomposition is a key process of nutrient and carbon cycling in terrestrial ecosystems. The decomposition process will likely be altered under ongoing climate change, both through direct effects on decomposer activity and through indirect effects caused by changes in litter quality. We studied how hydrological change indirectly affects decomposition via plant functional community restructuring caused by changes in plant species’ relative abundances (community‐weighted mean (CWM) traits and functional diversity). We further assessed how those indirect litter quality effects compare to direct effects. We set up a mesocosm experiment, in which sown grassland communities and natural turf pieces were subjected to different hydrological conditions (dryness and waterlogging) for two growing seasons. Species‐level mean traits were obtained from trait databases and combined with species’ relative abundances to assess functional community restructuring. We studied decomposition of mixed litter from these communities in a common “litterbed.” These indirect effects were compared to effects of different hydrological conditions on soil respiration and on decomposition of standard litter (direct effects). Dryness reduced biomass production in sown communities and natural turf pieces, while waterlogging only reduced biomass in sown communities. Hydrological stress caused profound shifts in species’ abundances and consequently in plant functional community composition. Hydrologically stressed communities had higher CMW leaf dry matter content, lower CMW leaf nitrogen content, and lower functional diversity. Lower CWM leaf N content and functional diversity were strongly related to slower decomposition. These indirect effects paralleled direct effects, but were larger and longer‐lasting. Species mean traits from trait databases had therefore considerable predictive power for decomposition. Our results show that stressful soil moisture conditions, that are likely to occur more frequently in the future, quickly shift species’ abundances. The resulting functional community restructuring will decelerate decomposition under hydrological stress.     

Conservation in the Context of Climate Change: Practical Guidelines for Land Protection at Local Scales

Climate change will affect the composition of plant and animal communities in many habitats and geographic settings. This presents a dilemma for conservation programs – will the portfolio of protected lands we now have achieve a goal of conserving biodiversity in the future when the ecological communities occurring within them change? Climate change will significantly alter many plant communities, but the geophysical underpinnings of these landscapes, such as landform, elevation, soil, and geological properties, will largely remain the same. Studies show that extant landscapes with a diversity of geophysical characteristics support diverse plant and animal communities. Therefore, geophysically diverse landscapes will likely support diverse species assemblages in the future, although which species and communities will be present is not altogether clear. Following protocols advanced in studies spanning large regions, we developed a down-scaled, high spatial resolution measure of geophysical complexity based on Ecological Land Units (ELUs) and examined the relationship between plant species richness, ecological community richness, and ELU richness (number of different ELU types). We found that extant landscapes with high ELU richness had a greater variety of ecological community types and high species richness of trees, shrubs, and herbaceous plants. We developed a spatial representation of diverse ELU landscapes to inform local conservation practitioners, such as land trusts, of potential conservation targets that will likely support diverse faunas and floras despite the impact of climate change.     

植物应对干旱胁迫的气孔调节

气孔是植物控制叶片与大气之间碳、水交换的重要门户,植物的生长和生存都依赖于叶片气孔对碳获取和水散失的调控.因此,气孔调节机理研究与气孔导度模型研发是精确模拟陆地生态系统碳、水循环过程不可或缺的内容.近年来,随着气候变化的加剧,干旱事件愈发频繁,对植物的存活、生长和分布产生深刻影响.为了深入理解植物碳-水耦合机理过程、预测全球变化下植物及群落的动态,开展植物应对干旱胁迫的气孔调节研究尤为重要.本文综述了植物在干旱胁迫条件下气孔调节机制和模型研究进展.首先阐述了植物气孔对干旱胁迫的主动调节与被动调节,讨论了气孔调节的演化过程,包括蕨类和石松类植物的被动水力调节、被子植物的主动调节和裸子植物的双重调节机制,认为裸子植物的气孔调节方式是植物进化过程中介于蕨类、石松类植物和被子植物之间的一种重要过渡类型.然后分析了气孔调节与水力调节的关系,讨论了“植物水势和气孔导度解耦”问题中存在的争议.之后介绍了基于水分利用效率假说和最大碳增益假说所建立的气孔导度优化模型的应用,并指出后者有更强的预测能力和应用前景.最后,为了有效减少植被对气候变化响应预测中的不确定性,提出了2个亟待开展的研究问题:将植物叶片的气孔调节功能研究由个体扩展到生态系统甚至更大尺度,改进陆地生态系统碳水循环机理模型;量化气孔调节的主动水力反馈过程,修正植物气孔功能水力模型.     

Assessing five decades of change in a high Arctic parasitoid community

Arctic ecosystems are fragile, and are particularly sensitive to the pressures of climate change. Both average temperature and precipitation have increased over the past five decades on Ellesmere Island, Nunavut, Canada in the high Arctic. Altered growth forms and increased biomass in dominant plant species on Ellesmere Island have been observed concurrent with the changing climate, but shifts in the diversity or rank abundance of plant and bird species have not been detected. Changes in diversity may take longer to appear, or may be assessed better using organisms with shorter generation times such as insects. We explored the ecological impacts of climate change on Ellesmere Island using historical and contemporary communities of ichneumonid wasps. We compared community diversity, functional composition, and body size of two common species using ichneumonid specimens collected in 1961–1965, 1980–1982, 1989–1992, and 2010. We found high compositional similarity between collections, overlapping estimates of species richness, no change in the proportion of idiobiont genera in the community, and no clear pattern in body size over time. The greatest amount of variation over time was detected in parasitoids of herbivores; proportionally fewer herbivore‐parasitizing genera were found in 2010 than in historical collections, and the two genera that were only found in one of the four collections were both parasitoids of herbivores. Our results point to the need to assess climate change effects in Arctic systems using a range of taxa, and responses to large‐scale environmental disturbances may be idiosyncratic and difficult to predict.     

Shedding light on shade: ecological perspectives of understorey plant life

Shade, in ecological sense, is not merely a lack of light, but a multi-faceted phenomenon that creates new and complex settings for community and ecosystem dynamics. Tolerating shade therefore affects plants’ ability to cope with other stressors, and also shape its interactions with surrounding organisms. The aim of this broad review was to map our current knowledge about how shade affects plants, plant communities and ecosystems – to gather together knowledge of what we know, but also to point out what we do not yet know. This review covers the following topics: the nature of shade, and ecological and physiological complexities related to growing under a canopy; plants’ capability of tolerating other stress factors while living under a shade – resource trade-offs and polytolerance of abiotic stress; ontogenetic effects of shade tolerance; coexistence patterns under the canopy – how shade determines the forest structure and diversity; shade-induced abiotic dynamics in understorey vegetation, including changing patterns of irradiance, temperature and humidity under the canopy; shade-driven plant–plant and plant–animal interactions – how shade mediates facilitation and stress, and how it creates differentiated environment for different herbivores and pollinators, including the role of volatile organic compounds. We also discuss the ways how vegetation in understorey environments will be affected by climate change, as shade might play a significant role in mitigating negative effects of climate change. Our review shows that living under a shade affects biotic and abiotic stress tolerance of plants, it also influences the outcomes of both symbiotic and competitive plant–plant and plant–animal interactions in a complex and dynamic manner. The current knowledge of shade-related mechanisms is rather ample, however there is much room for progress in integrating different implications of the multifaceted nature of shade into consistent and integral understanding how communities and ecosystems function.     

Soil heterogeneity buffers community response to climate change in species‐rich grassland

Climate change impacts on vegetation are mediated by soil processes that regulate rhizosphere water balance, nutrient dynamics, and ground‐level temperatures. For ecosystems characterized by high fine‐scale substrate heterogeneity such as grasslands on poorly developed soils, effects of climate change on plant communities may depend on substrate properties that vary at the scale of individuals (<m²), leading to fine‐scale shifts in community structure that may go undetected at larger scales. Here, we show in a long‐running climate experiment in species‐rich limestone grassland in Buxton, England (UK), that the resistance of the community to 15‐year manipulations of temperature and rainfall at the plot scale (9 m²) belies considerable community reorganization at the microsite (100 cm²) scale. In individual models of the abundance of the 25 most common species with respect to climate treatment and microsite soil depth, 13 species exhibited significant soil depth affinities, and nine of these have shifted their position along the depth gradient in response to one or more climate treatments. Estimates of species turnover across the depth gradient reviewed in relation to measurements of water potential, nitrogen supply, pH, and community biomass suggest that communities of shallow microsites are responding directly to microenvironmental changes induced by climate manipulation, while those of the deepest microsites are shifting in response to changes in competitive interference from more nutrient‐demanding species. Moreover, for several species in summer drought and winter heated treatments, climate response in deep microsites was opposite that of shallow microsites, suggesting microsite variation is contributing to community stability at the whole‐plot level. Our study thus demonstrates a strong link between community dynamics and substrate properties, and suggests ecosystems typified by fine‐scale substrate heterogeneity may possess a natural buffering capacity in the face of climate change.     

Plant species composition shifts in the Tatra Mts as a response to environmental change: a resurvey study after 90 years

Mountain vegetation is often considered highly sensitive to climate and land-use changes due to steep environmental gradients determining local plant species composition. In this study we present plant species compositional shifts in the Tatra Mts over the past 90 years and discuss the potential drivers of the changes observed. Using historical vegetation studies of the region from 1927, we resurveyed 76 vegetation plots, recording the vascular flora of each plot using the same methodology as in the original survey. We used an indirect method to quantify plant species compositional shifts and to indicate which environmental gradients could be responsible for these shifts: by calculating shifts in estimated species optima as reflected in shifts in the ecological indicator values of co-occurring species. To find shifts in species composition, focusing on each vegetation type separately, we used ordination (DCA). The species optimum changed significantly for at least one of the tested environmental gradients for 26 of the 95 plant species tested; most of these species changed in terms of the moisture indicator value. We found that the strongest shifts in species composition were in mylonite grassland, snowbed and hygrophilous tall herb communities. Changes in precipitation and increase in temperature were found to most likely drive compositional shifts in vegetation resurveyed. It is likely that the combined effect of climate change and cessation of sheep grazing has driven a species composition shift in granite grasslands communities.     

Animal behaviour shapes the ecological effects of ocean acidification and warming: moving from individual to community‐level responses

Biological communities are shaped by complex interactions between organisms and their environment as well as interactions with other species. Humans are rapidly changing the marine environment through increasing greenhouse gas emissions, resulting in ocean warming and acidification. The first response by animals to environmental change is predominantly through modification of their behaviour, which in turn affects species interactions and ecological processes. Yet, many climate change studies ignore animal behaviour. Furthermore, our current knowledge of how global change alters animal behaviour is mostly restricted to single species, life phases and stressors, leading to an incomplete view of how coinciding climate stressors can affect the ecological interactions that structure biological communities. Here, we first review studies on the effects of warming and acidification on the behaviour of marine animals. We demonstrate how pervasive the effects of global change are on a wide range of critical behaviours that determine the persistence of species and their success in ecological communities. We then evaluate several approaches to studying the ecological effects of warming and acidification, and identify knowledge gaps that need to be filled, to better understand how global change will affect marine populations and communities through altered animal behaviours. Our review provides a synthesis of the far‐reaching consequences that behavioural changes could have for marine ecosystems in a rapidly changing environment. Without considering the pervasive effects of climate change on animal behaviour we will limit our ability to forecast the impacts of ocean change and provide insights that can aid management strategies.     

Abrupt ecological changes in the last 800 years inferred from a mountainous bog using testate amoebae traits and multi-proxy data

Mountainous peatlands of Western Sudetes are considered a unique habitat in Central Europe. The region contains one of the largest raised bog complexes in temperate Europe and is thus of great importance for biodiversity conservation. In this first high-resolution study from this region we use long-term ecological data to assess how these mountain wetland ecosystems responded to anthropogenic impacts and climate change. We used testate amoebae morphological traits, micro- and macroscopic charcoal, pollen and plant macrofossils to reconstruct the history of peatland development over 800 years, illustrating shifts in its development and fire dynamics. Five hydrological stages of peatland development were recognized. Testate amoebae morphological traits reflected several abrupt ecological changes linked to anthropogenic modifications of landscape openness. A shift towards mixotrophic taxa, linked to hydrological change or shrubs expansion and shading, preceded aperture position change, which was associated to dust input through surrounding deforestation and simultaneous water-table increase. Fire reconstruction revealed increasing burning together with intensifying human activity, including the expansion of a nearby settlement. This study confirms the potential of testate amoeba communities and the use of morpho-functional traits as indicators of ecological effects of land-use change over long-temporal scales.     

干旱胁迫植物个体生理响应及其生态模型预测研究进展

全球气候突变导致干旱事件频发,进而易引发严重的植物衰退甚至死亡,聚焦植物尤其是树木死亡的生理学机制并期望基于此评估及预测气候变化导致植物死亡风险已成为热点话题。植物通过调整内在生理代谢过程,例如通过调节渗透物质的含量,来平衡渗透势、维持细胞膨压、调节植物激素的信号水平,诱导植物气孔开放程度降低,有利于植物保存水分、调控植物水通道蛋白的表达,进而保持体内水分稳定并对干旱胁迫做出快速响应。这些生理过程中的每一环调节都为了确保水分运输的效率和安全性,增加植物抗旱性以及生态系统稳定性。植物的抗旱性不仅体现在生理代谢方面的调节,还表现在植物水力特性与解剖结构间相辅相成。当植物改变水力特性的同时,其茎叶会在解剖结构上做出调整以满足植物在干旱环境下水分供需平衡,从而降低植物蒸腾水分散失、增强细胞储水并提高生存能力。植物应对水分胁迫的策略通常与水分消耗和碳获取之间的平衡有关,明晰植物水分消耗与光合碳获取间存在平衡关系的性状特征便于更好地理解植物的水分利用策略。然而,植物表现出的任意单一性状特征的强弱都无法代表整个植物适应逆境的优劣,未来只有通过将植物更多性状特征进行相互关联,以具有代表植物水力功能、结...