陆地生态系统野外增温控制实验的技术与方法

由于人类活动导致的碳排放急剧增加,工业革命以来全球地表温度显著增加约1℃,未来全球气候还将持续变暖,到21世纪末最高可升温4℃。这种前所未有的气候变化不仅影响陆地植被的适应策略,也深刻影响生态系统的结构和功能。其中陆地生态系统碳收支对全球变暖的反馈,是决定未来气候变化强度的关键因素,因此全球已经开展了大量的生态系统尺度的野外增温控制实验,研究生态系统碳收支对气温升高的响应,从而提高地球系统模型的预测精度。然而由于增温技术和方法的不同,不同研究的结果之间难以进行比较。该文系统总结了常见的野外增温技术和方法,包括主动增温和被动增温,阐述了其优缺点、适用对象以及相关研究成果。同时简要介绍了野外增温控制实验的前沿研究方向——新一代野外增温技术(包括全土壤剖面增温和全生态系统增温)和基于新一代增温技术开展的野外增温联网实验。     

全球变暖与陆地生态系统研究中的野外增温装置

由于化石燃料燃烧和森林砍伐等人类活动引起的地球大气层中温室气体(主要是二氧化碳)的富集已导致全球平均温度在20世纪升高了0.6 ℃,并将在本世纪继续上升1.4~5.8 ℃。这种地质历史上前所未有的全球变暖将对陆地植物和生态系统产生深远影响,并通过全球碳循环的改变反馈于全球气候变化。作为全球变化生态学的主要研究方法之一,生态系统增温实验能够为生态模型提供参数估计和模型验证。然而由于在世界各地使用的增温装置不同,使得各个生态系统之间的结果比较和整合难以实施,增加了模型预测的不确定性。该文通过比较几种常见的野外增温装置在模拟全球变暖情形时的优缺点,指出利用不同增温装置进行全球变暖研究中应注意的一些问题;同时探讨了全球变暖控制实验研究中的一些关键性的科学问题。     

增温对土壤有机碳矿化的影响研究综述

全球变暖的大背景下,土壤作为陆地生态系统中最大碳汇的载体,其微小变化都会引起大气CO2浓度显著的改变。土壤有机碳对气候变化的响应和适应对于预测未来气候变化具有十分重要的作用。然而,目前增温对土壤有机碳的影响及其影响机制仍存诸多未解决的问题。综述了目前土壤有机碳矿化的研究方式及增温对土壤有机碳矿化影响的国内外研究进展。结果发现增温往往会促进土壤有机碳排放,主要源于土壤微生物代谢活性或群落组成的改变。同时该排放强度因生态系统类型、增温方式和幅度以及增温季节和持续时间的不同而存在巨大差异,且长期增温反而使土壤微生物产生适应及驯化现象,从而降低或缓解陆地生态系统对全球变暖的正反馈效应。但这些结果大都基于温带实验,而原位增温实验对高生产力、多样性丰富的热带亚热带地区的影响是否与温带一致仍待进一步考证。室内模拟实验虽可深入研究温度对土壤有机碳矿化的影响机制,却无法真实反映野外自然环境。同时,野外增温方式及室内研究方式的多样均降低不同研究之间的可比性,进而难以预估由实验方法本身差异引起的结果变异。     

增温和放牧对草地土壤和生态系统呼吸的影响

草地生态系统作为世界陆地生态系统的主体类型,其土壤呼吸和生态系统呼吸是陆地生态系统碳循环的重要组成部分,土壤呼吸是未经扰动的土壤由于代谢活动而产生CO2的过程,生态系统呼吸包括地下部分的土壤呼吸和地上部分植被的自养呼吸。研究增温和放牧对草地土壤和生态系统呼吸的影响,可为预测未来气候变化条件下的全球碳收支以及草地的可持续经营与管理提供重要的科学依据。该文扼要综述了关于草地土壤和生态系统呼吸对增温和放牧的响应方面的研究。结果表明:草地土壤和生态系统呼吸对增温和放牧的响应非常复杂,受多种因素的综合影响,无论是增温还是放牧对草地土壤和生态系统呼吸的影响均具有不确定性,因草地类型、增温幅度、增温时间、放牧强度、放牧频度和放牧方式的不同而不同。在此基础上,指出了以后应加强研究的方向,草地的利用离不开放牧,对于未来气候变化条件下的草地,温度升高和放牧这两个因素必然是同时存在的,以前多数实验是单独研究增温或放牧对它们的影响,然而,这两者对草地生态系统的影响并非可加的,因此,需要加强增温与放牧的耦合试验,同时加强关于生态系统呼吸不同组分对两者的响应的研究,以便更好地理解增温和放牧的影响机制。另外,草地土壤和生态系统呼吸对增温和放牧的响应会随着时间的推移而发生变化,因而加强长期连续的试验观测很有必要。     

陆地生态系统凋落物分解对全球气候变暖的响应

陆地生态系统凋落物分解是全球碳收支的一个重要组成部分, 主要受气候、凋落物质量和土壤生物群落的综合控制。科学家们普遍认为全球气候变化将对陆地生态系统凋落物分解产生复杂而深远的影响。该文结合凋落物分解试验的常用方法——缩微试验、原位模拟实验和自然环境梯度实验, 归纳现有研究结果, 意在揭示全球气候变化对陆地生态系统凋落物分解的直接影响(温度对凋落物分解速率的影响)和间接影响(温度对凋落物质量、土壤微生物群落及植被型的影响)的普遍规律。各种研究方法都表明: 在水分条件理想的情况下, 温度升高往往能加快凋落物的分解速率; 原位模拟实验中, 凋落物分解速率因物种、增温方法和地理方位而异; 全球气候变化能改变凋落物质量, 但可能不会在短期内影响凋落物的分解速率; 凋落物质量和可分解性的种间差异远大于增温所引发的表型响应差异, 那么, 气候变化所引发的植物群落结构和物种组成的变化将对陆地生态系统凋落物分解产生更强烈的影响; 土壤生物群落如何响应全球气候变化, 进而怎样影响凋落物分解过程, 这些都还存在着极大的不确定性。     

区域尺度陆地生态系统碳收支及其循环过程研究进展

地球系统的碳库和碳循环过程变化是影响气候系统的重要因素,而陆地生态系统的碳收支及其循环过程机制研究一直是全球气候变化成因分析、变化趋势预测、减缓和适应对策分析领域的科学研究热点。回顾了过去几十年区域尺度陆地生态系统碳循环和碳收支研究领域的国际前沿及其关键科学问题,并分析了我国在该研究领域的科技需求和发展方向。当前国际科学研究的热点和前沿领域主要包括:生态系统和区域碳储量和碳收支的清查、综合计量与碳汇认证,陆地生态系统碳通量的联网观测及其循环过程机制,陆地生态系统碳循环过程对气候变化响应野外控制试验,陆地生态系统水、碳、氮循环及其耦合关系机制和模拟模型研究等,同时指出在这些研究领域依然存在且急需解决的关键科学问题。我国近期的科技工作重点工作应该是努力构建天-地-空一体化的碳储量和碳收支动态监测体系、开展生态系统碳-氮-水耦合循环及其区域调控管理的前瞻性研究,定量评价中国生态系统的碳收支状况和增汇潜力,评估各种典型生态系统增汇技术的经济效益,为国家尺度的温室气体管理和碳交易机制与政策体系的建立提供可报告、可度量和可核查的科学数据和技术支持。     

中国草地生态系统模拟增温实验的综合比较

近年来,全球增温对陆地生态系统的影响成为科学家研究的热点之一。草地生态系统是最重要的陆地生态系统之一,故研究草地生态系统对增温的响应尤为重要。本文通过对近10年国内外研究文献的回顾,论述了陆地生态系统增温实验的发展及其在中国草地生态系统的应用,从植物种群、群落、土壤、草地生态功能及动物等方面综合分析了增温对我国高寒草甸、亚高山草甸、羊草草甸、荒漠草原等不同草地类型的影响。结果表明,短期增温有利于草甸区禾本科植物的生长;在植物的不同生长时期增温对地上生物量的重要性存在差异;增温会改变植物的整体物候,显著降低羊草草甸土壤湿度;适度增温可以增加土壤微生物生物量碳、氮,促进高寒草甸土壤碳、磷循环。在此基础上,提出了将来应着重研究的几个科学问题。     

不同幅度的实验增温对藏北高寒草甸净生态系统碳交换的影响

全球气候变暖将对陆地生态系统(尤其是高寒草甸生态系统)碳循环产生深远影响。该研究依托中国科学院地理科学与资源研究所藏北高原草地生态系统研究站(那曲站), 设置不同增温幅度实验, 模拟未来2 ℃增温和4 ℃增温的情景, 探究不同增温幅度对青藏高原高寒草甸净生态系统碳交换(NEE)的影响。研究结果显示: 1)在2015年生长季(6-9月), 不增温和2 ℃增温处理下NEE小于0, 总体表现为碳汇, 而4 ℃增温处理下NEE大于0, 总体表现为碳源; 2)在生长季的6月、8月及整个生长季, 与不增温相比, 4 ℃增温处理显著提高了NEE, 而2 ℃增温处理没有显著改变NEE; 7月, 2 ℃和4 ℃增温处理均显著提高了NEE; 3)在半干旱的高寒草甸生态系统, 土壤水分是决定NEE的关键因素, 增温通过降低土壤水分而导致高寒草甸生态系统碳汇能力下降。该研究可为青藏高原高寒草甸生态系统应对未来气候变化提供基础数据和理论依据。     

森林土壤酶对环境变化的响应研究进展

全球气候变化已是不争的事实,对陆地生态系统特别是森林生态系统物质循环将产生显著的影响。土壤酶是森林土壤物质循环的主要限制因素之一,对气候变化的响应近年来受到广泛关注。由于森林土壤酶对全球气候变化的响应研究是预测未来环境变化对森林生态系统过程影响的关键,因此,着重综述了森林土壤酶对环境变化尤其是全球变暖和氮沉降响应方面的研究,并分析了未来研究的主要方向。环境变化会引起土壤p H、水分及其营养成分的变化,而这些变化会反作用于土壤酶的活性和稳定性。森林土壤酶对增温的响应,不仅与酶的种类以及增温的温度范围和持续时间有关,还与土壤类型有关,是多种因子综合作用的结果。森林土壤酶对氮添加的响应与林分类型和土层类型有关,受复合氮的影响更大。建议未来的研究应加强酶的基本性质对环境变化的响应研究,注重林分类型、土层类型导致的差异,强化多因素的交互作用,并进行长期、综合的观测。     

Experimental warming shifts coupling of carbon and nitrogen cycles in an alpine meadow

增温对高寒草甸生态系统碳氮循环耦合关系的影响 陆地生态系统碳吸收受土壤氮素可用性的调节。然而,全球变化背景下的不同生态系统组分的碳氮比及其所反映的碳氮循环耦合关系尚不十分清楚。本文运用数据同化的方法,将一个高寒草甸增温试验的14组数据同化到草地生态系统模型中,从而评估了增温如何影响陆地生态系统的碳氮循环耦合关系。研究结果表明,增温提高了土壤氮素的有效性,降低了土壤活性碳库的碳氮比,导致植物对土壤氮的吸收增加。但是由于植物叶片吸收的碳比吸收的氮增加更多,使得叶片中碳氮比增加,而根部的碳输入增加则低于氮的增加,导致根部的碳氮比减少。同时,增温降低了凋落物碳氮比,可能是在土壤高氮有效性的条件下,凋落物氮的固定得到增强;而且增温加速了凋落物的分解。同时增温还增加了慢速土壤有机质的碳氮比,使得该土壤碳库的碳固存潜力增大。由于大多数模型在不同的环境中通常使用相对固定的碳氮比,本研究所发现的气候变暖条件下碳氮比的差异变化可为模型参数化提供一个有效的参考,有利于模型对未来气候变化背景下生态系统碳氮耦合关系响应的预测。     

Response and adaptation of terrestrial ecosystem processes to climate warming

陆地生态系统包含一系列时空连续、尺度多元且互相联系的生态学过程。由于大部分生态学过程都受到温度调控, 因此气候变暖会对全球陆地生态系统产生深远的影响。近年来, 全球变化生态学的基本科学问题之一是陆地生态系统的关键过程如何响应与适应全球气候变暖。围绕该问题, 该文梳理了近年来的研究进展, 重点关注植物生理生态过程、物候期、群落动态、生产力及其分配、凋落物与土壤有机质分解、养分循环等过程对温度升高的响应与适应机理。通过定量分析近20年来发表于主流期刊的相关论文, 展望了该领域的前沿方向, 包括物种性状对生态系统过程的预测能力, 生物地球化学循环的耦合过程, 极端高温与低温事件的响应与适应机理, 不对称气候变暖的影响机理和基于过程的生态系统模拟预测等。基于这些研究进展, 该文建议进一步研究陆地生态系统如何适应气候变暖, 更多关注我国的特色生态系统类型, 并整合实验、观测或模型等研究手段开展跨尺度的合作研究。     

陆地生态系统过程对气候变暖的响应与适应

陆地生态系统包含一系列时空连续、尺度多元且互相联系的生态学过程。由于大部分生态学过程都受到温度调控, 因此气候变暖会对全球陆地生态系统产生深远的影响。近年来, 全球变化生态学的基本科学问题之一是陆地生态系统的关键过程如何响应与适应全球气候变暖。围绕该问题, 该文梳理了近年来的研究进展, 重点关注植物生理生态过程、物候期、群落动态、生产力及其分配、凋落物与土壤有机质分解、养分循环等过程对温度升高的响应与适应机理。通过定量分析近20年来发表于主流期刊的相关论文, 展望了该领域的前沿方向, 包括物种性状对生态系统过程的预测能力, 生物地球化学循环的耦合过程, 极端高温与低温事件的响应与适应机理, 不对称气候变暖的影响机理和基于过程的生态系统模拟预测等。基于这些研究进展, 该文建议进一步研究陆地生态系统如何适应气候变暖, 更多关注我国的特色生态系统类型, 并整合实验、观测或模型等研究手段开展跨尺度的合作研究。     

气候变暖对陆地生态系统碳循环的影响

作为全球变化的主要表现之一,气候变暖对全球陆地生态系统碳循环的影响巨大,揭示这一作用对于精确理解碳循环的过程和相关政策的制定具有重要的指导意义。该文综述了此领域近十几年来的主要研究工作,总结了陆地生态系统碳循环对气候变暖响应的主要内部机制及其过程,简述了相关模型的发展及其主要应用,并指出以往研究中存在的主要问题以及未来研究的主要方向。在气候变暖条件下,陆地生态系统碳循环的变化主要体现在以下几个方面:1)低纬度地区生态系统NPP一般表现为降低,而在中高纬度地区通常表现为增加,而在全球尺度上表现为NPP增加;2)土壤呼吸作用增强,但经过一段时间后表现出一定的适应性;3)高纬度地区的生态系统植被碳库表现为增加趋势,低纬度地区生态系统植被碳库变化不大,或略微降低,在全球尺度上表现为植被碳库增加;4)地表凋落物的产量和分解速率增加;5)土壤有机碳分解加速,进而减少土壤碳储存,同时植被碳库向土壤碳库的流动增加从而增加土壤碳库,这两种作用在不同生态系统的比重不同,在全球尺度上表现为土壤碳库的减少;6)尽管不同生态系统表现各异,总体上全球陆地生态系统表现为一个弱碳源。生物物理模型、生物地理模型和生物地球化学模型陆续被开发出来用于研究工作,并取得了一定的成果,但是研究结果仍然存在很大的不确定性。在未来的数年甚至是数十年间,气候变暖与全球变化的其它表现间的协同影响将是下一步的研究重点,气候变暖和陆地生态系统间的双向反馈作用机制是进行更准确研究的理论基础,生态系统结构和功能对气候变化的适应性是准确理解和预测未来气候情景下陆地生态系统碳循环的前提。     

Responses of terrestrial ecosystems to temperature and precipitation change: a meta‐analysis of experimental manipulation

Global mean temperature is predicted to increase by 2–7 °C and precipitation to change across the globe by the end of this century. To quantify climate effects on ecosystem processes, a number of climate change experiments have been established around the world in various ecosystems. Despite these efforts, general responses of terrestrial ecosystems to changes in temperature and precipitation, and especially to their combined effects, remain unclear. We used meta‐analysis to synthesize ecosystem‐level responses to warming, altered precipitation, and their combination. We focused on plant growth and ecosystem carbon (C) balance, including biomass, net primary production (NPP), respiration, net ecosystem exchange (NEE), and ecosystem photosynthesis, synthesizing results from 85 studies. We found that experimental warming and increased precipitation generally stimulated plant growth and ecosystem C fluxes, whereas decreased precipitation had the opposite effects. For example, warming significantly stimulated total NPP, increased ecosystem photosynthesis, and ecosystem respiration. Experimentally reduced precipitation suppressed aboveground NPP (ANPP) and NEE, whereas supplemental precipitation enhanced ANPP and NEE. Plant productivity and ecosystem C fluxes generally showed higher sensitivities to increased precipitation than to decreased precipitation. Interactive effects of warming and altered precipitation tended to be smaller than expected from additive, single‐factor effects, though low statistical power limits the strength of these conclusions. New experiments with combined temperature and precipitation manipulations are needed to conclusively determine the importance of temperature–precipitation interactions on the C balance of terrestrial ecosystems under future climate conditions.     

Effects of climate warming on carbon fluxes in grasslands— A global meta‐analysis

Climate warming will affect terrestrial ecosystems in many ways, and warming‐induced changes in terrestrial carbon (C) cycling could accelerate or slow future warming. So far, warming experiments have shown a wide range of C flux responses, across and within biome types. However, past meta‐analyses of C flux responses have lacked sufficient sample size to discern relative responses for a given biome type. For instance grasslands contribute greatly to global terrestrial C fluxes, and to date grassland warming experiments provide the opportunity to evaluate concurrent responses of both plant and soil C fluxes. Here, we compiled data from 70 sites (in total 622 observations) to evaluate the response of C fluxes to experimental warming across three grassland types (cold, temperate, and semi‐arid), warming methods, and short (≤3 years) and longer‐term (>3 years) experiment lengths. Overall, our meta‐analysis revealed that experimental warming stimulated C fluxes in grassland ecosystems with regard to both plant production (e.g., net primary productivity (NPP) 15.4%; aboveground NPP (ANPP) by 7.6%, belowground NPP (BNPP) by 11.6%) and soil respiration (Rs) (9.5%). However, the magnitude of C flux stimulation varied significantly across cold, temperate and semi‐arid grasslands, in that responses for most C fluxes were larger in cold than temperate or semi‐arid ecosystems. In semi‐arid and temperate grasslands, ecosystem respiration (Reco) was more sensitive to warming than gross primary productivity (GPP), while the opposite was observed for cold grasslands, where warming produced a net increase in whole‐ecosystem C storage. However, the stimulatory effect of warming on ANPP and Rs observed in short‐term studies (≤3 years) in both cold and temperate grasslands disappeared in longer‐term experiments (>3 years). These results highlight the importance of conducting long‐term warming experiments, and in examining responses across a wide range of climate.     

Effects of experimental warming on net ecosystem CO2 exchange in Northern Xizang alpine meadow

AimsGlobal warming could have profound effects on ecosystem carbon (C) fluxes in alpine ecosystems. The aim of our study is to examine the effects of gradient warming on net ecosystem carbon exchange (NEE).MethodsIn the Northern Tibetan Grassland Ecosystem Research Station (Nagqu station), Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Sciences, we conducted various levels of temperature increasing experiments (i.e., 2 °C and 4 °C increments). The warming was achieved using open-top chambers (OTCs). In total, there were three levels of temperature treatments (control, 2 °C and 4 °C increment), and four replicates for each treatment. The ecosystem NEE was monitored every five days during the growing season in 2015.Important findings Our findings highlight the importance of soil moisture in mediating the responses of NEE to climatic warming in alpine meadow ecosystem. The 4 °C warming significantly stimulated NEE,except for July measurements. The 2 °C warming had no effects on NEE during the growing season. Compared to the 2 °C warming, the 4 °C warming significantly stimulated NEE. The results showed that our targeted ecosystem acts as a carbon sink under 2 °C warming, whereas will act as a net carbon source under 4 °C warming in the future. This study provides basic data and theoretical basis for evaluating the alpine ecosystem’s responses to climate change.     

Soil warming alters nitrogen cycling in a New England forest: implications for ecosystem function and structure

Global climate change is expected to affect terrestrial ecosystems in a variety of ways. Some of the more well-studied effects include the biogeochemical feedbacks to the climate system that can either increase or decrease the atmospheric load of greenhouse gases such as carbon dioxide and nitrous oxide. Less well-studied are the effects of climate change on the linkages between soil and plant processes. Here, we report the effects of soil warming on these linkages observed in a large field manipulation of a deciduous forest in southern New England, USA, where soil was continuously warmed 5°C above ambient for 7 years. Over this period, we have observed significant changes to the nitrogen cycle that have the potential to affect tree species composition in the long term. Since the start of the experiment, we have documented a 45% average annual increase in net nitrogen mineralization and a three-fold increase in nitrification such that in years 5 through 7, 25% of the nitrogen mineralized is then nitrified. The warming-induced increase of available nitrogen resulted in increases in the foliar nitrogen content and the relative growth rate of trees in the warmed area. Acer rubrum (red maple) trees have responded the most after 7 years of warming, with the greatest increases in both foliar nitrogen content and relative growth rates. Our study suggests that considering species-specific responses to increases in nitrogen availability and changes in nitrogen form is important in predicting future forest composition and feedbacks to the climate system.     

A review on the effects of nitrogen and phosphorus addition on tree growth and productivity in forest ecosystems

Nitrogen (N) and phosphorus (P) inputs induced by anthropogenic activities and atmospheric N and P deposition have largely increased the availability of soil N and P in terrestrial ecosystems, which have considerably affected terrestrial carbon cycling processes. Tree growth and productivity in forest ecosystems play an important role in global carbon cycling, and determine the magnitude and direction of terrestrial carbon sequestration. Currently, a large number of field manipulation experiments have been conducted to investigate the effects of N and/or P addition on tree growth and forest productivity, but the results from these studies were inconsistent. Such inconsistent results might be affected by multiple factors, including biological, environmental and experimental variables. Here, we reviewed the present research status of the effects of N and P addition on tree growth and forest productivity in forest ecosystems based on three aspects, including the number of publications and experiments with field N and P addition, and the global distributions of these experiments. Then, we summarized the methods for assessing tree growth and forest productivity at ecosystem level in forest ecosystems, including relative growth rate and absolute increment. According to the related results, we reviewed the regulating factors that affect tree growth and productivity, and the potential mechanisms for such factors, including climate, tree size and stand age, plant functional traits (including type of tree-associated mycorrhizal fungi, N-fixation property of trees, and conservative and acquisitive functional traits), plant-microbe interaction, ambient nutrient (i.e., N and P) deposition rate, and experimental variables. Finally, we summarized the current studies, and pointed out five aspects that are urgently needed to provide further insights in future studies, including the physiological mechanism of how tree growth responds to N and P addition, the tradeoff and allocation among growth of various parts of tree under N and P addition, the role of plant functional traits in regulating and predicting the responses of tree growth to N and P addition, how the competition among trees regulates the responses of tree growth to N and P addition, and conducting long-term and coordinated distributed field experiments investigating the effects of N and P addition on tree growth and forest productivity at the global scale.