城市系统碳循环:特征、机理与理论框架

城市是地表受人类活动影响最深刻的区域,城市系统碳循环在全球和区域碳过程中具有重要的地位和作用.提出了城市“自然-社会”二元碳循环的概念,探讨了城市系统碳循环的一般特征;分析了城市系统碳循环的内部机理,主要包括:城市系统碳储量和碳输入/输出通量的主要过程和途径、城市系统碳储量、碳通量和碳流通的生命周期分析、城市系统碳输入和碳输出的类型划分等;提出了基于系统层次划分和碳流通过程的城市系统碳循环的研究框架,分析了城市自然系统和城市经济系统的主要碳流通过程和环节,构建了城市系统碳循环研究的思路和理论框架;最后提出了城市系统碳循环领域未来的研究重点.     

Reconciling Carbon-cycle Concepts, Terminology, and Methods

Recent projections of climatic change have focused a great deal of scientific and public attention on patterns of carbon (C) cycling as well as its controls, particularly the factors that determine whether an ecosystem is a net source or sink of atmospheric carbon dioxide (CO₂). Net ecosystem production (NEP), a central concept in C-cycling research, has been used by scientists to represent two different concepts. We propose that NEP be restricted to just one of its two original definitions—the imbalance between gross primary production (GPP) and ecosystem respiration (ER). We further propose that a new term—net ecosystem carbon balance (NECB)—be applied to the net rate of C accumulation in (or loss from [negative sign]) ecosystems. Net ecosystem carbon balance differs from NEP when C fluxes other than C fixation and respiration occur, or when inorganic C enters or leaves in dissolved form. These fluxes include the leaching loss or lateral transfer of C from the ecosystem; the emission of volatile organic C, methane, and carbon monoxide; and the release of soot and CO₂ from fire. Carbon fluxes in addition to NEP are particularly important determinants of NECB over long time scales. However, even over short time scales, they are important in ecosystems such as streams, estuaries, wetlands, and cities. Recent technological advances have led to a diversity of approaches to the measurement of C fluxes at different temporal and spatial scales. These approaches frequently capture different components of NEP or NECB and can therefore be compared across scales only by carefully specifying the fluxes included in the measurements. By explicitly identifying the fluxes that comprise NECB and other components of the C cycle, such as net ecosystem exchange (NEE) and net biome production (NBP), we can provide a less ambiguous framework for understanding and communicating recent changes in the global C cycle.     

Trait diversity shapes the carbon cycle

Trait evolution is shaped by carbon economics at the organismal level. Here, we expand this idea to the ecosystem level and show how the trait diversity of ecological communities influences the carbon cycle. Systematic shifts in trait diversity will likely trigger changes in the carbon cycle.     

Negative feedback processes following drainage slow down permafrost degradation

The sustainability of the vast Arctic permafrost carbon pool under climate change is of paramount importance for global climate trajectories. Accurate climate change forecasts, therefore, depend on a reliable representation of mechanisms governing Arctic carbon cycle processes, but this task is complicated by the complex interaction of multiple controls on Arctic ecosystem changes, linked through both positive and negative feedbacks. As a primary example, predicted Arctic warming can be substantially influenced by shifts in hydrologic regimes, linked to, for example, altered precipitation patterns or changes in topography following permafrost degradation. This study presents observational evidence how severe drainage, a scenario that may affect large Arctic areas with ice‐rich permafrost soils under future climate change, affects biogeochemical and biogeophysical processes within an Arctic floodplain. Our in situ data demonstrate reduced carbon losses and transfer of sensible heat to the atmosphere, and effects linked to drainage‐induced long‐term shifts in vegetation communities and soil thermal regimes largely counterbalanced the immediate drainage impact. Moreover, higher surface albedo in combination with low thermal conductivity cooled the permafrost soils. Accordingly, long‐term drainage effects linked to warming‐induced permafrost degradation hold the potential to alleviate positive feedbacks between permafrost carbon and Arctic warming, and to slow down permafrost degradation. Self‐stabilizing effects associated with ecosystem disturbance such as these drainage impacts are a key factor for predicting future feedbacks between Arctic permafrost and climate change, and, thus, neglect of these mechanisms will exaggerate the impacts of Arctic change on future global climate projections.     

Spatial early warning signals for impending regime shifts: A practical framework for application in real‐world landscapes

Prediction of ecosystem response to global environmental change is a pressing scientific challenge of major societal relevance. Many ecosystems display nonlinear responses to environmental change, and may even undergo practically irreversible ‘regime shifts’ that initiate ecosystem collapse. Recently, early warning signals based on spatiotemporal metrics have been proposed for the identification of impending regime shifts. The rapidly increasing availability of remotely sensed data provides excellent opportunities to apply such model‐based spatial early warning signals in the real world, to assess ecosystem resilience and identify impending regime shifts induced by global change. Such information would allow land‐managers and policy makers to interfere and avoid catastrophic shifts, but also to induce regime shifts that move ecosystems to a desired state. Here, we show that the application of spatial early warning signals in real‐world landscapes presents unique and unexpected challenges, and may result in misleading conclusions when employed without careful consideration of the spatial data and processes at hand. We identify key practical and theoretical issues and provide guidelines for applying spatial early warning signals in heterogeneous, real‐world landscapes based on literature review and examples from real‐world data. Major identified issues include (1) spatial heterogeneity in real‐world landscapes may enhance reversibility of regime shifts and boost landscape‐level resilience to environmental change (2) ecosystem states are often difficult to define, while these definitions have great impact on spatial early warning signals and (3) spatial environmental variability and socio‐economic factors may affect spatial patterns, spatial early warning signals and associated regime shift predictions. We propose a novel framework, shifting from an ecosystem perspective towards a landscape approach. The framework can be used to identify conditions under which resilience assessment with spatial remotely sensed data may be successful, to support well‐informed application of spatial early warning signals, and to improve predictions of ecosystem responses to global environmental change.     

The impact of projected increases in urbanization on ecosystem services

Alteration in land use is likely to be a major driver of changes in the distribution of ecosystem services before 2050. In Europe, urbanization will probably be the main cause of land-use change. This increase in urbanization will result in spatial shifts in both supplies of ecosystem services and the beneficiaries of those services; the net outcome of such shifts remains to be determined. Here, we model changes in urban land cover in Britain based on large (16%) projected increases in the human population by 2031, and the consequences for three different services--flood mitigation, agricultural production and carbon storage. We show that under a scenario of densification of urban areas, the combined effect of increasing population and loss of permeable surfaces is likely to result in 1.7 million people living within 1 km of rivers with at least 10 per cent increases in projected peak flows, but that increasing suburban 'sprawl' will have little effect on flood mitigation services. Conversely, losses of stored carbon and agricultural production are over three times as high under the sprawl as under the 'densification' urban growth scenarios. Our results illustrate the challenges of meeting, but also of predicting, future demands and patterns of ecosystem services in the face of increasing urbanization.     

大尺度森林碳循环过程模拟模型综述

森林生态系统碳循环是全球陆地生态系统碳循环的重要组成部分,而碳循环模型已经成为研究森林碳循环的必要手段。森林碳循环模型可以分为统计模型和过程模型,其中过程模型以其完整的理论框架、严谨的结构分析和清晰的过程机理,逐渐占据了主导地位。从地球化学过程模型、陆面物理过程模型和生物过程模型等3个方面综述区域尺度到全球尺度(本文称为大尺度)森林碳循环过程模型研究进展,论述了各类模型的主要特征、优缺点以及应用现状,探讨了森林碳循环模拟研究中存在的问题,并讨论了森林碳循环过程模型的主流研究方向。可为不同空间尺度下森林生态系统碳循环模拟模型的选择提供参考,以及为森林碳循环研究提供借鉴。     

Commentary: Carbon Metabolism of the Terrestrial Biosphere: A Multitechnique Approach for Improved Understanding

Understanding terrestrial carbon metabolism is critical because terrestrial ecosystems play a major role in the global carbon cycle. Furthermore, humans have severely disrupted the carbon cycle in ways that will alter the climate system and directly affect terrestrial metabolism. Changes in terrestrial metabolism may well be as important an indicator of global change as the changing temperature signal. Improving our understanding of the carbon cycle at various spatial and temporal scales will require the integration of multiple, complementary and independent methods that are used by different research communities. Tools such as air sampling networks, inverse numerical methods, and satellite data (top-down approaches) allow us to study the strength and location of the global- and continental-scale carbon sources and sinks. Bottom-up studies provide estimates of carbon fluxes at finer spatial scales and examine the mechanisms that control fluxes at the ecosystem, landscape, and regional scales. Bottom-up approaches include comparative and process studies (for example, ecosystem manipulative experiments) that provide the necessary mechanistic information to develop and validate terrestrial biospheric models. An iteration and reiteration of top-down and bottom-up approaches will be necessary to help constrain measurements at various scales. We propose a major international effort to coordinate and lead research programs of global scope of the carbon cycle. Received 7 May 1999; accepted 28 September 1999.     

Underestimated ecosystem carbon turnover time and sequestration under the steady state assumption: A perspective from long‐term data assimilation

It is critical to accurately estimate carbon (C) turnover time as it dominates the uncertainty in ecosystem C sinks and their response to future climate change. In the absence of direct observations of ecosystem C losses, C turnover times are commonly estimated under the steady state assumption (SSA), which has been applied across a large range of temporal and spatial scales including many at which the validity of the assumption is likely to be violated. However, the errors associated with improperly applying SSA to estimate C turnover time and its covariance with climate as well as ecosystem C sequestrations have yet to be fully quantified. Here, we developed a novel model‐data fusion framework and systematically analyzed the SSA‐induced biases using time‐series data collected from 10 permanent forest plots in the eastern China monsoon region. The results showed that (a) the SSA significantly underestimated mean turnover times (MTTs) by 29%, thereby leading to a 4.83‐fold underestimation of the net ecosystem productivity (NEP) in these forest ecosystems, a major C sink globally; (b) the SSA‐induced bias in MTT and NEP correlates negatively with forest age, which provides a significant caveat for applying the SSA to young‐aged ecosystems; and (c) the sensitivity of MTT to temperature and precipitation was 22% and 42% lower, respectively, under the SSA. Thus, under the expected climate change, spatiotemporal changes in MTT are likely to be underestimated, thereby resulting in large errors in the variability of predicted global NEP. With the development of observation technology and the accumulation of spatiotemporal data, we suggest estimating MTTs at the disequilibrium state via long‐term data assimilation, thereby effectively reducing the uncertainty in ecosystem C sequestration estimations and providing a better understanding of regional or global C cycle dynamics and C‐climate feedback.     

陆地碳循环的动态非平衡假说

生态系统维持物质与能量的动态平衡是地球系统孕育与维持生物多样性的重要基础。自工业革命以来, 人类活动导致陆地生态系统的碳循环转变为动态非平衡,进而使陆地生态系统的结构与功能出现许多难以预测的变化动态。本文阐释了陆地生态系统碳循环的动态非平衡假说。该假说构建于陆地碳循环内部过程的四点基本特征和五类外部驱动因素。基于这些内部特征与外部因素, 本文归纳了陆地生态系统碳循环动态非平衡在不同时间与空间尺度的表达现象, 并从观测、实验与模型的角度讨论了其检测方法。陆地生态系统碳循环的动态非平衡假说不仅有助于我们理解复杂的陆地碳循环现象, 也为预测未来陆地碳汇动态提供了新的理论框架。     

海岛陆地生态系统固碳估算方法

陆地生态系统在调节全球碳平衡和减缓全球气候变化中起着重要作用。海岛作为一种特殊的生态系统,生物群落和环境与大陆基本相似。虽然海岛生态结构相对简单,物种的丰富程度比大陆低,但对全球碳循环也有一定的影响。在海岛陆地生态系统中,森林和灌草的种属相对较少,且不同纬度的海岛森林植被种属差异明显,可采用典型样地清查和生物量模型估算相结合的方法估算乔木层和灌草层的碳储量。采用模型估算固碳潜力时,根据海岛生态环境的特殊性,综合考虑岛陆面积、季节、风向、坡度、坡向、海拔、平均温度、降雨量、土壤理化性质等参数对其碳储量估算的影响。海岛植被生物多样性影响其土壤碳储存的生态服务功能,利用多元统计分析方法,建立岛陆植物物种丰度与土壤碳储量的空间回归模型,明确植物多样性的改变对岛陆土壤固碳能力的影响。此外,从土壤固碳的角度而言,海岛土壤-植物-微生物间相互作用是其重要的研究方向。利用现代分子生物学技术,研究海岛陆地生态系统的土壤-植物-微生物相互作用关系,有利于海岛土壤固碳潜力估算精度的提高。     

稳定性同位素技术与Keeling曲线法在陆地生态系统碳/水交换研究中的应用

稳定性同位素技术和Keeling曲线法是现代生态学研究的重要手段和方法之一。稳定性同位素能够整合生态系统复杂的生物学、生态学和生物地球化学过程在时间和空间尺度上对环境变化的响应。Keeling曲线法是以生物过程前后物质平衡理论为基础,将CO₂或H₂O的同位素组成(δD、δ¹³C或δ¹⁸O)与其对应浓度测量结合起来,将生态系统净碳通量区分为光合固定和呼吸释放通量,或将整个生态系统水分蒸散区分为植物蒸腾和土壤蒸发。在全球尺度上,稳定性同位素技术、Keeling曲线法与全球尺度陆地生态系统模型相结合,还可区分陆地生态系统和海洋生态系统对全球碳通量的贡献以及不同植被类型(C₃或C₄)在全球CO₂同化量中所占的比例。然而,生态系统的异质性使得稳定性同位素技术和Keeling曲线法从冠层尺度外推到生态系统、区域或全球尺度时存在有一定程度的不确定性。此外,取样时间、地点的选取也会影响最终的研究结果。尽管如此,随着分析手段的不断精确和研究方法的日趋完善,稳定性同位素技术和Keeling曲线法与其它测量方法(如微气象法)的有机结合将成为未来陆地生态系统碳/水交换研究的重要手段和方法之一。     

Agricultural modifications of hydrological flows create ecological surprises

Agricultural expansion and intensification have altered the quantity and quality of global water flows. Research suggests that these changes have increased the risk of catastrophic ecosystem regime shifts. We identify and review evidence for agriculture-related regime shifts in three parts of the hydrological cycle: interactions between agriculture and aquatic systems, agriculture and soil, and agriculture and the atmosphere. We describe the processes that shape these regime shifts and the scales at which they operate. As global demands for agriculture and water continue to grow, it is increasingly urgent for ecologists to develop new ways of anticipating, analyzing and managing nonlinear changes across scales in human-dominated landscapes.     

森林生态系统碳循环动态仿真系统的设计

模型方法是森林碳循环研究的有力工具.在Simulink环境下设计开发了通用的森林生态系统碳循环动态仿真系统FORCASS,从仿真系统的模式框架、设计方案和开发过程方面进行综合分析表明,FORCASS具有可行性.该仿真系统具有如下特点：1）将森林生态系统划分为植被碳库、枯落物碳库、土壤碳库和动物碳库4个分室,考虑了众多碳流转移项,具有较高的机理性和解释性;2）仿真系统基于过程,以植被器官生物量碳储量Richards生长方程为驱动项,带入差分方程组进行计算,可操作性高,能够实现林龄变化下的植被净第一性生产力（NPP）、净生态系统生产力（NEP）等多种输出;3）仿真系统基于通用的碳循环模式框架建立,可扩展性能良好.     

Shifting plant phenology in response to global change

Plants are finely tuned to the seasonality of their environment, and shifts in the timing of plant activity (i.e. phenology) provide some of the most compelling evidence that species and ecosystems are being influenced by global environmental change. Researchers across disciplines have observed shifting phenology at multiple scales, including earlier spring flowering in individual plants and an earlier spring green-up' of the land surface revealed in satellite images. Experimental and modeling approaches have sought to identify the mechanisms causing these shifts, as well as to make predictions regarding the consequences. Here, we discuss recent advances in several fields that have enabled scaling between species responses to recent climatic changes and shifts in ecosystem productivity, with implications for global carbon cycling.     

Human‐aided admixture may fuel ecosystem transformation during biological invasions: theoretical and experimental evidence

Biological invasions can transform our understanding of how the interplay of historical isolation and contemporary (human‐aided) dispersal affects the structure of intraspecific diversity in functional traits, and in turn, how changes in functional traits affect other scales of biological organization such as communities and ecosystems. Because biological invasions frequently involve the admixture of previously isolated lineages as a result of human‐aided dispersal, studies of invasive populations can reveal how admixture results in novel genotypes and shifts in functional trait variation within populations. Further, because invasive species can be ecosystem engineers within invaded ecosystems, admixture‐induced shifts in the functional traits of invaders can affect the composition of native biodiversity and alter the flow of resources through the system. Thus, invasions represent promising yet under‐investigated examples of how the effects of short‐term evolutionary changes can cascade across biological scales of diversity. Here, we propose a conceptual framework that admixture between divergent source populations during biological invasions can reorganize the genetic variation underlying key functional traits, leading to shifts in the mean and variance of functional traits within invasive populations. Changes in the mean or variance of key traits can initiate new ecological feedback mechanisms that result in a critical transition from a native ecosystem to a novel invasive ecosystem. We illustrate the application of this framework with reference to a well‐studied plant model system in invasion biology and show how a combination of quantitative genetic experiments, functional trait studies, whole ecosystem field studies and modeling can be used to explore the dynamics predicted to trigger these critical transitions.     

Trends in tuna carbon isotopes suggest global changes in pelagic phytoplankton communities

Considerable uncertainty remains over how increasing atmospheric CO₂ and anthropogenic climate changes are affecting open‐ocean marine ecosystems from phytoplankton to top predators. Biological time series data are thus urgently needed for the world's oceans. Here, we use the carbon stable isotope composition of tuna to provide a first insight into the existence of global trends in complex ecosystem dynamics and changes in the oceanic carbon cycle. From 2000 to 2015, considerable declines in δ¹³C values of 0.8‰–2.5‰ were observed across three tuna species sampled globally, with more substantial changes in the Pacific Ocean compared to the Atlantic and Indian Oceans. Tuna recorded not only the Suess effect, that is, fossil fuel‐derived and isotopically light carbon being incorporated into marine ecosystems, but also recorded profound changes at the base of marine food webs. We suggest a global shift in phytoplankton community structure, for example, a reduction in ¹³C‐rich phytoplankton such as diatoms, and/or a change in phytoplankton physiology during this period, although this does not rule out other concomitant changes at higher levels in the food webs. Our study establishes tuna δ¹³C values as a candidate essential ocean variable to assess complex ecosystem responses to climate change at regional to global scales and over decadal timescales. Finally, this time series will be invaluable in calibrating and validating global earth system models to project changes in marine biota.     

Fire in Australian savannas: from leaf to landscape

Savanna ecosystems comprise 22% of the global terrestrial surface and 25% of Australia (almost 1.9 million km²) and provide significant ecosystem services through carbon and water cycles and the maintenance of biodiversity. The current structure, composition and distribution of Australian savannas have coevolved with fire, yet remain driven by the dynamic constraints of their bioclimatic niche. Fire in Australian savannas influences both the biophysical and biogeochemical processes at multiple scales from leaf to landscape. Here, we present the latest emission estimates from Australian savanna biomass burning and their contribution to global greenhouse gas budgets. We then review our understanding of the impacts of fire on ecosystem function and local surface water and heat balances, which in turn influence regional climate. We show how savanna fires are coupled to the global climate through the carbon cycle and fire regimes. We present new research that climate change is likely to alter the structure and function of savannas through shifts in moisture availability and increases in atmospheric carbon dioxide, in turn altering fire regimes with further feedbacks to climate. We explore opportunities to reduce net greenhouse gas emissions from savanna ecosystems through changes in savanna fire management.     

Human-induced salinity changes impact marine organisms and ecosystems

Climate change is fundamentally altering marine and coastal ecosystems on a global scale. While the effects of ocean warming and acidification on ecology and ecosystem functions and services are being comprehensively researched, less attention is directed toward understanding the impacts of human-driven ocean salinity changes. The global water cycle operates through water fluxes expressed as precipitation, evaporation, and freshwater runoff from land. Changes to these in turn modulate ocean salinity and shape the marine and coastal environment by affecting ocean currents, stratification, oxygen saturation, and sea level rise. Besides the direct impact on ocean physical processes, salinity changes impact ocean biological functions with the ecophysiological consequences are being poorly understood. This is surprising as salinity changes may impact diversity, ecosystem and habitat structure loss, and community shifts including trophic cascades. Climate model future projections (of end of the century salinity changes) indicate magnitudes that lead to modification of open ocean plankton community structure and habitat suitability of coral reef communities. Such salinity changes are also capable of affecting the diversity and metabolic capacity of coastal microorganisms and impairing the photosynthetic capacity of (coastal and open ocean) phytoplankton, macroalgae, and seagrass, with downstream ramifications on global biogeochemical cycling. The scarcity of comprehensive salinity data in dynamic coastal regions warrants additional attention. Such datasets are crucial to quantify salinity-based ecosystem function relationships and project such changes that ultimately link into carbon sequestration and freshwater as well as food availability to human populations around the globe. It is critical to integrate vigorous high-quality salinity data with interacting key environmental parameters (e.g., temperature, nutrients, oxygen) for a comprehensive understanding of anthropogenically induced marine changes and its impact on human health and the global economy.     

阴山北麓农牧交错区退耕地草地生态系统碳交换及水分利用效率

内蒙古阴山北麓农牧交错区由于长期不合理的开垦造成了荒漠草地生态系统碳交换等生态功能的显著丧失。我国20世纪末开始实施的退耕还林还草工程产生了大量退耕地,随着自然恢复演替,这些退耕地的生态功能得到了有效的修复,其巨大的碳汇潜力成为了荒漠草地生态系统碳循环研究的热点。研究通过空间代替时间的方法,对内蒙古阴山北麓典型区域——武川县周边无干扰的荒漠草原以及3个退耕恢复阶段草地的生态系统CO₂交换(NEE、GEP和ER)、水分利用效率(WUE)以及生物量等指标进行了实地测量。结果表明：(1)随着退耕恢复演替时间的推移,生态系统CO₂交换呈显著上升趋势,演替晚期植被NEE与未受干扰的荒漠草地无明显差异;(2)生态系统水分利用效率的变化趋势与生态系统CO₂交换基本一致,但已退耕17年后的退耕地WUE仍没有恢复至未受干扰荒漠草地的水平;(3)导致以上结果的原因主要与退耕地地上植被生物量的恢复以及一、二年生植物和多年生植物比例的演替变化有关。结果表明荒漠草地退耕地恢复过程中生态系统功能的恢复可能并非是同时的,而是分阶段有选择进行的。