植物群落向土壤有机碳输入及其对气候变暖的响应研究进展

植物群落作为陆地生态系统土壤有机碳的主要来源,可通过地表凋落物分解、细根周转和根系分泌物等方式将光合作用同化的碳输入到土壤中。全球气候变暖正深刻地影响植物群落的分布、结构与功能,改变森林地上和地下凋落物产量与分解速率和根系分泌过程,从而改变植物群落向土壤输入有机碳数量。本文综述了植物群落向土壤有机碳输入过程及其对气候变暖的响应研究进展。研究表明,气候变暖可通过影响植物群落生产直接影响凋落物产量和根系分泌过程,还可通过改变凋落物分解环境条件、凋落物基质质量和分解者群落结构与活性等非生物与生物因子而间接作用于凋落物向土壤有机碳输入过程。气候变暖还可通过影响植物根系性状、根系分泌物化学组成等间接影响植物根系向土壤输入的碳量,但其具体机制还需深入探讨。未来的研究应该关注气候变暖导致植物群落结构改变进而影响土壤有机碳输入的具体机制以及粗木质残体对土壤有机碳输入的贡献,同时还应注重植物凋落物与根系分泌过程的整合研究,以期更全面地认识气候变暖背景下植物群落对土壤碳库及碳循环过程的贡献。     

全球气候变暖对凋落物分解的影响

凋落物分解作为生态系统核心过程,参与生态系统碳的周转与循环,影响生态系统碳的收支平衡,调控生态系统对全球气候变暖的反馈结果。全球气候变暖通过环境因素、凋落物数量和质量以及分解者3个方面,直接或间接地作用于凋落物分解过程,并进一步影响土壤养分周转和碳库动态。气候变暖可通过升高温度和改变实际蒸散量等环境因素直接作用于凋落物分解。气候变暖可引起植物物种短期内碳、氮和木质素等化学性质的改变以及群落中物种组成的长期变化从而改变凋落物质量。在凋落物分解过程中,土壤分解者亚系统作为主要生命组分(土壤动物和微生物)彼此相互作用、相互协调共同参与调节凋落物的分解过程。凋落物分解可以通过改变土壤微生物量、微生物活动和群落结构来加快微生物养分的固定或矿化,以形成新的养分利用模式来改变土壤有机质从而对气候变化做出响应。未来凋落物分解的研究方向应基于大尺度跨区域分解实验和长期实验,关注多个因子交互影响下,分解过程中碳、氮养分释放、地上/地下凋落物分解生物学过程与联系、分解者亚系统营养级联效应等方面。     

陆地生态系统地下碳输入与输出过程研究进展

生态系统地下碳输入与输出过程是陆地生态系统碳分配和转化的核心,并直接影响着全球碳循环。陆地生态系统凋落物、根系周转、根系分泌物、土壤有机碳、土壤微生物和土壤呼吸是地下碳输入与输出过程中的重要组成部分。由于这些组分非常复杂且其研究技术和方法受到限制,目前人们对陆地生态系统地下碳输入与输出过程尚缺乏全面的认识,故在陆地生态系统碳循环研究中存在诸多的不确定性。该文概述了凋落物、根系周转、根系分泌物、土壤有机碳、土壤微生物和土壤呼吸的研究方法,以及它们对气候变化的响应,探讨了陆地生态系统地下碳输入与输出过程中的研究难点,并对未来需要深入探究的一些领域进行了展望。     

改变碳输入对沂蒙山区典型次生林土壤微生物碳源代谢功能的影响

凋落物和根系向土壤的碳输入是森林生态系统的关键过程,输入量及组分的变化直接影响森林土壤碳汇功能和生产力。在沂蒙山区栎类天然次生林中,开展添加/去除凋落物及去除根系的定位控制试验。于控制试验开展21个月后,采用Biolog Eco微平板培养法,研究凋落物和根系对土壤微生物碳源代谢功能的影响。结果表明,凋落物倍增处理增加了土壤微生物碳源代谢功能,增加了对糖类和胺类的代谢能力。去除凋落物处理、去除根系处理和无输入处理都降低了土壤微生物碳源代谢功能。去除凋落物处理降低土壤微生物碳源代谢功能的幅度大于去除根系处理,表明当前条件下凋落物对土壤微生物碳源代谢功能的影响大于根系,但如果抛除掉去除根系处理中残留根系的影响,凋落物和根系对土壤微生物碳源代谢功能的相对大小可能会发生变化。土壤有机碳含量、铵态氮含量显著影响微生物碳源代谢多样性（P<0.05）,并与碳源代谢功能正相关。凋落物倍增处理通过增加土壤铵态氮和有机碳含量,增加微生物碳源代谢功能,去除凋落物处理和无输入处理通过降低土壤铵态氮和有机碳含量,降低微生物碳源代谢功能。结果深化了碳输入途径（地上凋落物与地下根系）和数量（凋落物倍增、凋落物去除与对照）对温带栎类天然次生林土壤碳代谢过程的认识。     

冻融季温带森林土壤真菌群落结构及功能类群对不同土壤有机碳输入的响应

根系与凋落物有机碳输入变化对土壤生物群落的影响研究是目前学术界关注的热点问题,但冻融季不同有机碳输入方式将对土壤真菌群落结构及功能类群产生何种影响尚不明确。土壤真菌群落是调节森林生态系统稳定性的重要因素,有助于维持生态系统生产力时间尺度的稳定性。为了探索冻融季温带森林土壤真菌群落对控制根系和凋落物有机碳输入方式的响应特征,通过在帽儿山生态站设置4种碳源输入控制处理植物残体添加去除(DIRT):去除凋落物仅根系输入处理、去除根系仅凋落物输入处理、无碳源输入处理和同时进行根系与凋落物输入处理,采用ITS rDNA高通量测序技术和FUNGuild功能预测平台,来分析控制根系和凋落物有机碳输入方式对温带森林土壤真菌群落结构和功能类群的影响。研究结果显示：(1)不同有机碳输入方式改变了土壤真菌类群的多度：与自然生长状态下有机碳输入方式相比,根系有机碳输入比凋落物有机碳输入对土壤真菌类群多度影响更明显,去除根系碳源输入处理使真菌群落中子囊菌门含量升高19.52%,担子菌门含量下降16.77%。(2)有机碳输入方式对土壤真菌群落功能类群产生影响：与自然生长状态下有机碳输入方式相比,去除根系碳源输入处...     

草地植物根系碳储量和碳流转对CO2浓度升高的响应

植物根系是陆地生态系统重要的碳汇和矿质养分库,也是土壤中碳及养分的主要来源,只有深入认识CO2浓度升高下根系的碳汇功能和根系周转对土壤碳库的影响,才能准确预测生态系统对全球变化的响应与反馈调节作用.介绍了CO2浓度升高对草地植物根系生物量、根系凋落物的数量和品质以及根系周转速率的影响,指出研究植物体内碳向根分配格局的变化趋势必须考虑CO2浓度升高的直接和间接两方面作用;在预测根系碳库储量的动态变化时,需要区分不同功能根系组分的生物量;为更准确估算根系周转速率,有必要确立草地植物根系直径与其寿命之间的关系;CO2浓度升高普遍提高根系凋落物的C/N,但以此判定其在土壤中的分解速率快慢并不可靠,需要进一步从机理上探究根系凋落物分解的控制因素.     

添加混合凋落物对沙丘草地土壤有机碳矿化的影响

土壤有机碳矿化是调节温室气体排放、土壤有机质形成以及土壤生物和植物营养供应的重要过程,植物残体分解释放CO_2对土壤有机碳矿化有着重要影响。通过对科尔沁沙地沙丘草地4种优势植物叶凋落物的混合培养试验,测定了凋落物培养过程中CO_2释放速率及其累积释放量,比较了混合凋落物CO_2释放量实测值与预测值的差异,分析了凋落物化学成分和物种多样性(包括物种丰富度和物种组成)与土壤有机碳矿化的相关关系,以期解释添加混合凋落物对土壤有机碳矿化的影响。结果表明,混合凋落物物种丰富度对土壤有机碳矿化的影响不显著,而凋落物一些化学性质与土壤有机碳矿化紧密相关;所有混合组合处理中,80%的凋落物组合处理对土壤有机碳矿化产生显著(P0.05)的非加和效应;氮含量较高的豆科植物达乌里胡枝子凋落物与禾本科植物凋落物混合后土壤有机碳矿化表现极显著(P0.001)的协同非加和效应,而禾本科植物凋落物交互混合后土壤有机碳矿化产生显著拮抗非加和效应,这可能是凋落物化学成分相似或凋落物叶片的空间异质性引起的。     

胶州湾滨海湿地凋落物分解对土壤有机碳矿化的影响

通过室内培养试验和三维荧光光谱技术(3D-EEMs),开展了胶州湾滨海湿地碱蓬、芦苇和互花米草的分解对土壤有机碳矿化的影响研究。结果表明,凋落物的添加提高了土壤有机碳矿化速率和累积矿化量,具体表现为碱蓬互花米草芦苇空白对照;乘幂曲线模型能较好地描述有机碳矿化速率和累积矿化量的变化趋势。光谱分析表明,分解过程中类蛋白荧光强度始终大于类腐殖酸荧光强度;利用荧光区域积分进行定量分析表明,不同处理下类蛋白质物质占比最高,类腐殖质物质次之;荧光参数表明,土壤有机碳的芳香化程度在培养期间先升高后降低。凋落物分解通过增加土壤中的营养物质,提高了土壤中微生物活性,从而改变了土壤有机碳的结构和化学组分,且凋落物的植被特征决定了其对土壤有机碳矿化影响程度的大小。     

凋落物输入对三江平原弃耕农田土壤基础呼吸和活性碳组分的影响

土壤有机碳的积累主要由土壤有机质的输入与输出间的净平衡决定的,植被的恢复和凋落物质的大量输入是土壤恢复的先决条件,凋落物的输入在土壤恢复过程中起着至关重要的作用.通过对不同类型凋落物输入到三江平原弃耕农田后土壤的基础呼吸、溶解有机碳(DOC)和土壤微生物量碳(MBC)的研究表明:相同种类凋落物输入后,输入到土壤总有机碳(TOC)背景值低的凋落物被微生物降解的速率大于TOC背景值高的土壤,TOC较低的土壤能够加快微生物对输入凋落物的分解,不利于有机质的积累;不同类型凋落物的输入使土壤基础呼吸、DOC和MBC等活性组分的生成和降解产生差异,改变了凋落物的降解速率,在三江平原研究的4种主要植被类型中,人工林凋落物最容易降解,小叶章、大豆的降解能力次之,玉米是最难降解的凋落物.     

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。     

Transplantation of organic surface horizons of boreal soils into warmer regions alters microbiology but not the temperature sensitivity of decomposition

Changes in soil carbon, the largest terrestrial carbon pool, are critical for the global carbon cycle, atmospheric CO₂ levels and climate. Climate warming is predicted to be most pronounced in the northern regions and therefore the large soil carbon pool residing in boreal forests will be subject to larger global warming impact than soil carbon pools in the temperate or the tropical forest. A major uncertainty in current estimates of the terrestrial carbon balance is related to decomposition of soil organic matter (SOM). We hypothesized that when soils are exposed to warmer climate the structure of the ground vegetation will change much more rapidly than the dominant tree species. This change will alter the quality and amount of litter input to the soil and induce changes in microbial communities, thus possibly altering the temperature sensitivity of SOM decomposition. We transferred organic surface soil sections from the northern borders of the boreal forest zone to corresponding forest sites in the southern borders of the boreal forest zone and studied the effects of warmer climate after an adaptation period of 2 years. The results showed that initially ground vegetation and soil microbial community structure and community functions were different in northern and southern forest sites and that 2 years of exposure to warmer climate was long enough to cause changes in these ecological indicators. The rate of SOM decomposition was approximately equally sensitive to temperature irrespective of changes in vegetation or microbial communities in the studied forest sites. However, as temperature sensitivity of the decomposition increases with decreasing temperature regime, the proportional increase in the decomposition rate in northern latitudes could lead to significant carbon losses from the soils.     

Decoupling the direct and indirect effects of climate on plant litter decomposition: Accounting for stress‐induced modifications in plant chemistry

Decomposition of plant litter is a fundamental ecosystem process that can act as a feedback to climate change by simultaneously influencing both the productivity of ecosystems and the flux of carbon dioxide from the soil. The influence of climate on decomposition from a postsenescence perspective is relatively well known; in particular, climate is known to regulate the rate of litter decomposition via its direct influence on the reaction kinetics and microbial physiology on processes downstream of tissue senescence. Climate can alter plant metabolism during the formative stage of tissues and could shape the final chemical composition of plant litter that is available for decomposition, and thus indirectly influence decomposition; however, these indirect effects are relatively poorly understood. Climatic stress disrupts cellular homeostasis in plants and results in the reprogramming of primary and secondary metabolic pathways, which leads to changes in the quantity, composition, and organization of small molecules and recalcitrant heteropolymers, including lignins, tannins, suberins, and cuticle within the plant tissue matrix. Furthermore, by regulating metabolism during tissue senescence, climate influences the resorption of nutrients from senescing tissues. Thus, the final chemical composition of plant litter that forms the substrate of decomposition is a combined product of presenescence physiological processes through the production and resorption of metabolites. The changes in quantity, composition, and localization of the molecular construct of the litter could enhance or hinder tissue decomposition and soil nutrient cycling by altering the recalcitrance of the lignocellulose matrix, the composition of microbial communities, and the activity of microbial exo‐enzymes via various complexation reactions. Also, the climate‐induced changes in the molecular composition of litter could differentially influence litter decomposition and soil nutrient cycling. Compared with temperate ecosystems, the indirect effects of climate on litter decomposition in the tropics are not well understood, which underscores the need to conduct additional studies in tropical biomes. We also emphasize the need to focus on how climatic stress affects the root chemistry as roots contribute significantly to biogeochemical cycling, and on utilizing more robust analytical approaches to capture the molecular composition of tissue matrix that fuel microbial metabolism.     

Litter decomposition in grasslands of Central North America (US Great Plains)

One of the major concerns about global warming is the potential for an increase in decomposition and soil respiration rates, increasing CO₂ emissions and creating a positive feedback between global warming and soil respiration. This is particularly important in ecosystems with large belowground biomass, such as grasslands where over 90% of the carbon is allocated belowground. A better understanding of the relative influence of climate and litter quality on litter decomposition is needed to predict these changes accurately in grasslands. The Long‐Term Intersite Decomposition Experiment Team (LIDET) dataset was used to evaluate the influence of climatic variables (temperature, precipitation, actual evapotranspiration, and climate decomposition index), and litter quality (lignin content, carbon : nitrogen, and lignin : nitrogen ratios) on leaf and root decomposition in the US Great Plains. Wooden dowels were used to provide a homogeneous litter quality to evaluate the relative importance of above and belowground environments on decomposition. Contrary to expectations, temperature did not explain variation in root and leaf decomposition, whereas precipitation partially explained variation in root decomposition. Percent lignin was the best predictor of leaf and root decomposition. It also explained most variation in root decomposition in models which combined litter quality and climatic variables. Despite the lack of relationship between temperature and root decomposition, temperature could indirectly affect root decomposition through decreased litter quality and increased water deficits. These results suggest that carbon flux from root decomposition in grasslands would increase, as result of increasing temperature, only if precipitation is not limiting. However, where precipitation is limiting, increased temperature would decrease root decomposition, thus likely increasing carbon storage in grasslands. Under homogeneous litter quality, belowground decomposition was faster than aboveground and was best predicted by mean annual precipitation, which also suggests that the high moisture in soil accelerates decomposition belowground.     

Biodiversity and ecosystem productivity: implications for carbon storage

Recent experiments have found that Net Primary Productivity (NPP) can often be a positive saturating function of plant species and functional diversity. These findings raised the possibility that more diverse ecosystems might store more carbon as a result of increased photosynthetic inputs. However, carbon inputs will not only remain in plant biomass, but will be translocated to the soil via root exudation, fine root turnover, and litter fall. Thus, we must consider not just plant productivity (NPP), but also net productivity of the whole ecosystem (NEP), which itself measures net carbon storage. We currently know little about how plant diversity could influence soil processes that return carbon back to the atmosphere, such as heterotrophic respiration and decomposition of organic matter. Nevertheless, it is clear that any effects on such processes could make NPP a poor predictor of whole-ecosystem productivity, and potentially the ability of the ecosystem to store carbon. We examine the range of mechanisms by which plant diversity could influence net ecosystem productivity, incorporating processes involved with carbon uptake (productivity), loss (autotrophic and heterotrophic respiration), and residence time within the system (decomposition rate). Understanding the relationship between plant diversity and ecosystem carbon dynamics must be made a research priority if we wish to provide information relevant to global carbon policy decisions. This goal is entirely feasible if we utilize some basic methods for measuring the major fluxes of carbon into and out of the ecosystem.     

Disentangling the Litter Quality and Soil Microbial Contribution to Leaf and Fine Root Litter Decomposition Responses to Reduced Rainfall

Climate change-induced rainfall reductions in Mediterranean forests negatively affect the decomposition of plant litter through decreased soil moisture. However, the indirect effects of reduced precipitation on litter decomposition through changes in litter quality and soil microbial communities are poorly studied. This is especially the case for fine root litter, which contributes importantly to forests plant biomass. Here we analyzed the effects of long-term (11 years) rainfall exclusion (29% reduction) on leaf and fine root litter quality, soil microbial biomass, and microbial community-level physiological profiles in a Mediterranean holm oak forest. Additionally, we reciprocally transplanted soils and litter among the control and reduced rainfall treatments in the laboratory, and analyzed litter decomposition and its responses to a simulated extreme drought event. The decreased soil microbial biomass and altered physiological profiles with reduced rainfall promoted lower fine root—but not leaf—litter decomposition. Both leaf and root litter, from the reduced rainfall treatment, decomposed faster than those from the control treatment. The impact of the extreme drought event on fine root litter decomposition was higher in soils from the control treatment compared to soils subjected to long-term rainfall exclusion. Our results suggest contrasting mechanisms driving drought indirect effects on above-(for example, changes in litter quality) and belowground (for example, shifts in soil microbial community) litter decomposition, even within a single tree species. Quantifying the contribution of these mechanisms relative to the direct soil moisture-effect is critical for an accurate integration of litter decomposition into ecosystem carbon dynamics in Mediterranean forests under climate change.     

湿地枯落物分解及其对全球变化的响应

综述了当前湿地枯落物分解及其对全球变化响应的研究动态。湿地枯落物分解研究已随研究方法的改进而不断深化;当前湿地枯落物分解过程研究主要集中在有机质组分和元素含量变化特征的探讨上;湿地枯落物分解同时受生物因素（即枯落物性质以及参与分解的异养微生物和土壤动物的种类、数量和活性等）和非生物因素（即枯落物分解过程的外部环境条件,包括气候条件、水分条件、酸碱度与盐分条件以及湿地沉积的行为与特征等）的制约;模型已成为湿地枯落物分解研究的重要手段,对其研究也在不断深化。还讨论了湿地枯落物分解对于全球变化的响应,指出全球变暖、大气CO2浓度上升、干湿沉降及其化学组成改变可能对枯落物分解产生的直接、间接和综合影响。最后,指出了当前该领域研究尚存在的问题以及今后亟需加强的几个研究方面。     

Partitioning the effect of composition and diversity of tree communities on leaf litter decomposition and soil respiration

The decomposition of plant material is an important ecosystem process influencing both carbon cycling and soil nutrient availability. Quantifying how plant diversity affects decomposition is thus crucial for predicting the effect of the global decline in plant diversity on ecosystem functioning. Plant diversity could affect the decomposition process both directly through the diversity of the litter, and/or indirectly through the diversity of the host plant community and its affect on the decomposition environment. Using a biodiversity experiment with trees in which both functional and taxonomic diversity were explicitly manipulated independently, we tested the effects of the functional diversity and identity of the living trees separately and in combination with the functional diversity and identity of the decomposing litter on rates of litter decomposition and soil respiration. Plant traits, predominantly leaf chemical and physical traits, were correlated with both litter decomposition and soil respiration rates. Surface litter decomposition, quantified by mass loss in litterbags, was best explained by abundance‐weighted mean trait values of tree species from which the litter was assembled (functional identity). In contrast, soil respiration, which includes decomposition of dissolved organic carbon and root respiration, was best explained by the variance in trait values of the host trees (functional diversity). This research provides insight into the effect of loss of tree diversity in forests on soil processes. Such understanding is essential to predicting changes in the global carbon budget brought on by biodiversity loss.     

外来植物入侵对陆地生态系统地下碳循环及碳库的影响

生物入侵是当今全球性重大环境问题之一, 是全球变化的主要研究内容.评价外来植物入侵对于生态系统影响的研究多集中在地上部分,对于生态系统地下部分影响的研究相对较少.陆地生态系统地下部分对于生态系统过程的重要性之一体现在它处于生态系统碳分配过程的核心环节.入侵种通过影响群落凋落物的输入数量、质量以及输入时间,影响到对于土壤的碳输入,而入侵种与土著种根系的差异以及入侵种对微生物群落的影响是造成土壤呼吸强度发生变化的主要因素,前者土壤呼吸强度一般比后者高.多数研究表明外来植物入侵对生态系统地下碳循环和碳库产生影响,但由于入侵植物种类较多以及研究地点环境条件的不同,关于外来植物入侵对于土壤碳库和土壤有机碳矿化影响的研究结论并不统一.最后,提出了今后该研究领域应加强的一些建议和方向.     

Disentangling direct and indirect effects of water table drawdown on above‐ and belowground plant litter decomposition: consequences for accumulation of organic matter in boreal peatlands

Pristine peatlands are carbon (C)‐accumulating wetland ecosystems sustained by a high water table (WT) and consequent anoxia that slows down decomposition. Persistent WT drawdown as a response to climate and/or land‐use change affects decomposition either directly through environmental factors such as increased oxygenation, or indirectly through changes in plant community composition. This study attempts to disentangle the direct and indirect effects of WT drawdown by measuring the relative importance of environmental parameters (WT depth, temperature, soil chemistry) and litter type and/or litter chemical quality on the 2‐year decomposition rates of above‐ and belowground litter (altogether 39 litter types). Consequences for organic matter accumulation were estimated based on the annual litter production. The study sites were chosen to form a three‐stage chronosequence from pristine (undrained) to short‐term (years) and long‐term (decades) WT drawdown conditions at three nutrient regimes. The direct effects of WT drawdown were overruled by the indirect effects through changes in litter type composition and production. Short‐term responses to WT drawdown were small. In long‐term, dramatically increased litter inputs resulted in large accumulation of organic matter in spite of increased decomposition rates. Furthermore, the quality of the accumulated matter greatly changed from that accumulated in pristine conditions. Our results show that the shift in vegetation composition as a response to climate and/or land‐use change is the main factor affecting peatland ecosystem C cycle, and thus dynamic vegetation is a necessity in any model applied for estimating responses of C fluxes to changing environment. We provide possible grouping of litter types into plant functional types that the models could utilize. Furthermore, our results clearly show a drop in soil summer temperature as a response to WT drawdown when an initially open peatland converts into a forest ecosystem, which has not yet been considered in the existing models.