Root exudates and their ecological consequences in forest ecosystems: Problems and perspective北大核心CSCD

Researches on rhizosphere ecological processes and the underlying mechanisms have become one of the most active and sensitive hotspots in soil science. Root exudates have specialized roles in mediating the nutrient cycling and signal transduction within root-soil-microbe interactions. They are the key driving factors in regulating the functions of rhizosphere micro-ecosystem, and serve as a major premise for the concept and ecological processes in rhizosphere. However, due to the instinctive advantages of crops, such as short life cycles and convenient operation, most previous studies on root exudation mainly focused on agricultural ecosystems and were primarily targeted at providing practical guidelines. In contrast, there have been relatively few investigations on root exudates of trees, which highly limited the comprehensive knowledge of the potential mechanisms of root exudates in mediating soil biogeochemical processes in forest ecosystems. Hence, in this review, based on the main findings in our previous studies and the emerging frontiers in rhizosphere ecology, we specifically reviewed the ecological consequences and key remaining challenges in researches on root exudation in forests. Finally, we identify several topics and research outlooks for guiding future work to facilitate studies on root exudation and its ecological consequences in forest ecosystems. © Chinese Journal of Plant Ecology     

Advances and perspective in research on plant-soil-microbe interactions mediated by root exudates

根系分泌物是植物与土壤进行物质交换和信息传递的重要载体物质, 是植物响应外界胁迫的重要途径, 是构成植物不同根际微生态特征的关键因素, 也是根际对话的主要调控者。根系分泌物对于生物地球化学循环、根际生态过程调控、植物生长发育等均具有重要功能, 尤其是在调控根际微生态系统结构与功能方面发挥着重要作用, 调节着植物-植物、植物-微生物、微生物-微生物间复杂的互作过程。植物化感作用、作物间套作、生物修复、生物入侵等都是现代农业生态学的研究热点, 它们都涉及十分复杂的根际生物学过程。越来越多的研究表明, 不论是同种植物还是不同种植物之间相互作用的正效应或是负效应, 都是由根系分泌物介导下的植物与特异微生物共同作用的结果。近年来, 随着现代生物技术的不断完善, 有关土壤这一“黑箱”的研究方法与技术取得了长足的进步, 尤其是各种宏组学技术(meta-omics technology), 如环境宏基因组学、宏转录组学、宏蛋白组学、宏代谢组学等的问世, 极大地推进了人们对土壤生物世界的认知, 尤其是对植物地下部生物多样性和功能多样性的深层次剖析, 根际生物学特性的研究成果被广泛运用于指导生产实践。深入系统地研究根系分泌物介导下的植物-土壤-微生物的相互作用方式与机理, 对揭示土壤微生态系统功能、定向调控植物根际生物学过程、促进农业生产可持续发展等具有重要的指导意义。该文综述了根系分泌物的概念、组成及功能, 论述了根系分泌物介导下植物与细菌、真菌、土壤动物群之间的密切关系, 总结了探索根际生物学特性的各种研究技术及其优缺点, 并对该领域未来的研究方向进行了展望。     

根系分泌物收集技术研究进展

根系分泌物在调控陆地生态系统根际微环境间的物质、能量和信息交流中具有重要作用.构建准确、适用的根系分泌物收集方法,对根系分泌物的种类、含量及其对环境变化的响应等信息的精准获取是理解植物根系-土壤界面生态过程与信息交流的关键环节和前提.目前,传统或新型的根系分泌物收集技术都致力于认知根系分泌物中化合物种类多样性和含量变化,但根系的生长高度依赖周围介质和生长环境,使根系分泌物的收集很难避开对根系本身的损伤、土壤颗粒对根系分泌物的吸附和释放,以及微生物代谢等因素的干扰,导致不同根系分泌物收集方法都存在各自的优缺点.本研究从室内收集和野外原位收集两方面系统综述了当前应用较广的一些传统和新型根系分泌物收集技术,并总结和比较了每种收集方法的优缺点;在此基础上,基于森林根际生态学过程野外原位研究的重要性和代表性,结合当前根系分泌物研究的不足,展望了未来森林根系分泌物野外原位收集技术构建中值得关注的3个重点方向,旨在为相关研究者开展根系分泌物收集与作用研究提供参考.     

根分泌物对根际矿物营养及根际微生物的效应

综述了根系分泌物对植物生长的生理生态学效应,并就根系分泌物的定义、产生机制、组成成分和影响因素等方面进行了讨论。指出根系分泌物在缓解低矿物营养胁迫对植株造成的伤害及决定根际微生物的种群密度和数量方面起着重要的作用;根系分泌物的产生机制多样,组成成分复杂,影响因素繁多。对根分泌物的深入研究有助于进一步了解植物体与土壤间进行的生理生化过程及其调控机制。     

根分泌物及其生态效应

根分泌物是个古老而年轻的研究领域。早在18,19世纪,人们(Plenk,1795;Decandolle,1830)就观察到根分泌物对邻近植株的促生和抑制作用,1904年Hilter提出“根际”的概念,标志着人们对根分泌物及其生态效应的进一步认识。此后人们对根分泌物研究逐步展开。Lyon和Willson(1920)发现,生长于无菌水培液中的植物能释放有机物,为深入研究奠定了基础。但很长一段时间这个研究领域一     

The role of root exudates and allelochemicals in the rhizosphere

Plant roots serve a multitude of functions in the plant including anchorage, provision of nutrients and water, and production of exudates with growth regulatory properties. The root–soil interface, or rhizosphere, is the site of greatest activity within the soil matrix. Within this matrix, roots affect soil structure, aeration and biological activity as they are the major source of organic inputs into the rhizosphere, and are also responsible for depletion of large supplies of inorganic compounds. Roots are very complicated morphologically and physiologically, and their metabolites are often released in large quantities into the soil rhizosphere from living root hairs or fibrous root systems. Root exudates containing root-specific metabolites have critical ecological impacts on soil macro and microbiota as well as on the whole plant itself. Through the exudation of a wide variety of compounds, roots impact the soil microbial community in their immediate vicinity, influence resistance to pests, support beneficial symbioses, alter the chemical and physical properties of the soil, and inhibit the growth of competing plant species. In this review, we outline recent research on root exudation and the role of allelochemicals in the rhizosphere by studying the case of three plants that have been shown to produce allelopathic root exudates: black walnut, wheat and sorghum     

植物根系分泌物主要生态功能研究进展

根系分泌物在植物根系-土壤-微生物互作过程及其生态反馈机制中发挥重要作用。在植物根际复杂网络互作过程中, 根系分泌物被认为是“根际对话”的媒介, 其在调控植物适应微生境、缓解根际养分竞争及构建根际微生物群落结构方面意义重大。该文结合国内外该领域主要研究成果, 综述了根系分泌物对植物生长、土壤微生物特性及土壤养分循环的影响, 并展望了未来根系分泌物的研究方向。     

Research Advances in the Main Ecological Functions of Root Exudates

根系分泌物在植物根系-土壤-微生物互作过程及其生态反馈机制中发挥重要作用。在植物根际复杂网络互作过程中, 根系分泌物被认为是“根际对话”的媒介, 其在调控植物适应微生境、缓解根际养分竞争及构建根际微生物群落结构方面意义重大。该文结合国内外该领域主要研究成果, 综述了根系分泌物对植物生长、土壤微生物特性及土壤养分循环的影响, 并展望了未来根系分泌物的研究方向。     

N添加对西南亚高山红桦林根系分泌物及其介导的养分转化过程的影响

目前缺乏对根系分泌物通量以及相关生态后果对不同氮（N）沉降水平响应方向和幅度的深入理解,该研究以西南亚高山典型的红桦（Betula albosinensis）林为研究对象,通过野外原位N添加试验模拟不同氮沉降水平（对照组,0 kg?hm^-2?a^-1;低氮处理,25 kg?hm^-2?a^-1;高氮处理,50 kg?hm^-2?a^-1）,分析了红桦林根系分泌物C输入通量及其介导的根际土壤养分循环过程对不同N添加水平的差异化响应,试图揭示不同N添加处理对红桦根系分泌物C输入通量及其介导的土壤养分转化过程的影响。结果表明：（1）N添加显著抑制了红桦林根系分泌物C输入速率（其中低氮（N25）条件下单位根生物量根系分泌速率均值降低约14.87%）和年C输入通量（低氮条件下降低了约45.01%）（P<0.05）,其高氮处理的抑制效应更强。（2）N添加显著抑制了红桦林N矿化速率及其相关的微生物胞外酶活性（P<0.05）,并显著降低了其根际效应;N沉降显著抑制了根系分泌物C输入通量及其介导的土壤养分转化过程,并且这种抑制效应随N沉降水平的升高而增强。该研究结果可丰富全球气候变化下森林地下碳养分循环过程的认识和理解。     

根系分泌物介导下植物-土壤-微生物互作关系研究进展与展望

根系分泌物是植物与土壤进行物质交换和信息传递的重要载体物质, 是植物响应外界胁迫的重要途径, 是构成植物不同根际微生态特征的关键因素, 也是根际对话的主要调控者。根系分泌物对于生物地球化学循环、根际生态过程调控、植物生长发育等均具有重要功能, 尤其是在调控根际微生态系统结构与功能方面发挥着重要作用, 调节着植物-植物、植物-微生物、微生物-微生物间复杂的互作过程。植物化感作用、作物间套作、生物修复、生物入侵等都是现代农业生态学的研究热点, 它们都涉及十分复杂的根际生物学过程。越来越多的研究表明, 不论是同种植物还是不同种植物之间相互作用的正效应或是负效应, 都是由根系分泌物介导下的植物与特异微生物共同作用的结果。近年来, 随着现代生物技术的不断完善, 有关土壤这一“黑箱”的研究方法与技术取得了长足的进步, 尤其是各种宏组学技术(meta-omics technology), 如环境宏基因组学、宏转录组学、宏蛋白组学、宏代谢组学等的问世, 极大地推进了人们对土壤生物世界的认知, 尤其是对植物地下部生物多样性和功能多样性的深层次剖析, 根际生物学特性的研究成果被广泛运用于指导生产实践。深入系统地研究根系分泌物介导下的植物-土壤-微生物的相互作用方式与机理, 对揭示土壤微生态系统功能、定向调控植物根际生物学过程、促进农业生产可持续发展等具有重要的指导意义。该文综述了根系分泌物的概念、组成及功能, 论述了根系分泌物介导下植物与细菌、真菌、土壤动物群之间的密切关系, 总结了探索根际生物学特性的各种研究技术及其优缺点, 并对该领域未来的研究方向进行了展望。     

Regulation and function of root exudates

Root-secreted chemicals mediate multi-partite interactions in the rhizosphere, where plant roots continually respond to and alter their immediate environment. Increasing evidence suggests that root exudates initiate and modulate dialogue between roots and soil microbes. For example, root exudates serve as signals that initiate symbiosis with rhizobia and mycorrhizal fungi. In addition, root exudates maintain and support a highly specific diversity of microbes in the rhizosphere of a given particular plant species, thus suggesting a close evolutionary link. In this review, we focus mainly on compiling the information available on the regulation and mechanisms of root exudation processes, and provide some ideas related to the evolutionary role of root exudates in shaping soil microbial communities.     

Effects of nitrogen enrichment on root exudative carbon inputs in Sibiraea angustata shrubbery at the eastern fringe of Qinghai-Xizang Plateau

Aims Understanding the responses of root exudative carbon (C) to increasing nitrogen deposition is important for predicting carbon cycling in terrestrial ecosystems. However, fewer studies have investigated the dynamics of root exudation in shrubbery ecosystems compared to forests and grassland ecosystems. This objective of this study was to determine the effects of nitrogen fertilization on the rate and C flux of root exudates.Methods Three levels of nitrogen addition treatments were applied to a Sibiraea angustata shrubbery ecosystem situated at the eastern fringe of Qinghai-Xizang Plateau, including N₀ (without nitrogen application), N₅ (nitrogen addition rate of 5 g·m^-2·a^-1), and N₁₀ (nitrogen addition rate of 10 g·m^-2·a^-1), respectively, in 5 m ´ 5 m plots. Root exudates were collected in June, August and October of 2015, using a modified culture-based cuvette system. Root biomass in each plot was measured with root core method.Important findings The rates of root exudates on biomass, length, and surface area basis all displayed apparent seasonal variations during the experimental period, with the magnitude ranked in the order of: August > June > October, consistent with changes in soil temperature at 5 cm depth. With increases in the nitrogen addition rate, the rate of root exudates on biomass, length, and area basis all trended lower. Compared with the control (N₀), the N₅ and N₁₀ treatments significantly reduced fine root biomass in the Sibiraea angustata shrubbery, by 23.36% and 33.84%, respectively. The decreasing root exudation and fine root biomass in response to nitrogen addition significantly decreased C flux of root exudates. Our results provide additional evidences toward a robust theoretical foundation for better understanding soil C-nutrient cycling process mediated by root exudation inputs in Alpine shrubbery ecosystems under various environmental changes.     

Plant growth strategy determines the magnitude and direction of drought-induced changes in root exudates in subtropical forests

Root exudates are an important pathway for plant–microbial interactions and are highly sensitive to climate change. However, how extreme drought affects root exudates and the main components, as well as species-specific differences in response magnitude and direction, are poorly understood. In this study, root exudation rates of total carbon (C) and its components (e.g., sugar, organic acid, and amino acid) were measured under the control and extreme drought treatments (i.e., 70% throughfall reduction) by in situ collection of four tree species with different growth rates in a subtropical forest. We also quantified soil properties, root morphological traits, and mycorrhizal infection rates to examine the driving factors underlying variations in root exudation. Our results showed that extreme drought significantly decreased root exudation rates of total C, sugar, and amino acid by 17.8%, 30.8%, and 35.0%, respectively, but increased root exudation rate of organic acid by 38.6%, which were largely associated with drought-induced changes in tree growth rates, root morphological traits, and mycorrhizal infection rates. Specifically, trees with relatively high growth rates were more responsive to drought for root exudation rates compared with those with relatively low growth rates, which were closely related to root morphological traits and mycorrhizal infection rates. These findings highlight the importance of plant growth strategy in mediating drought-induced changes in root exudation rates. The coordinations among root exudation rates, root morphological traits, and mycorrhizal symbioses in response to drought could be incorporated into land surface models to improve the prediction of climate change impacts on rhizosphere C dynamics in forest ecosystems.     

Relevance of Rhizosphere Research to the Ecological Risk Assessment of Trace Metals in Soils

The chemical, mineralogical, and microbial properties of the rhizosphere of a range of forested ecosystems were studied to identify the key processes controlling the distribution and fate of trace metals at the soil–root interface. The results of our research indicate that: (1) the rhizosphere is a soil microenvironment where properties (e.g., pH, organic matter, microbes) and processes (nutrient and water absorption, exudation) differ markedly from those of the adjacent bulk soil; (2) the rhizosphere is a corrosive medium where the weathering and neoformation of soil solid phases are enhanced; (3) the concentrations of solid-phase and water-soluble trace metals like Cd, Cu, Ni, Pb, and Zn are generally higher in the rhizosphere as shown by both macroscopic and microscopic approaches; (4) a larger fraction of water-soluble metals is complexed by dissolved organic substances in the rhizosphere; and (5) soil microorganisms play, either directly or indirectly, a distinct role on metal speciation, in particular Cu and Zn, in the rhizosphere. These results improve our capacity to estimate metal speciation and bioavailability at the soil–root interface. Furthermore, the research emphasizes the crucial physical position occupied by the rhizosphere with respect to the process of elemental uptake by plants and its key functional role in the transfer of trace metals along the food chain. We conclude that the properties and processes of the rhizosphere should be viewed as key components of assessments of the ecological risks associated with the presence of trace metals in soils.     

A review of rhizosphere carbon flow modelling

Rhizosphere processes play a key role in nutrient cycling in terrestrial ecosystems. Plant rhizodeposits supply low-molecular weight carbon substrates to the soil microbial community, resulting in elevated levels of activity surrounding the root. Mechanistic compartmental models that aim to model carbon flux through the rhizosphere have been reviewed and areas of future research necessary to better calibrate model parameters have been identified. Incorporating the effect of variation in bacterial biomass physiology on carbon flux presents a considerable challenge to experimentalists and modellers alike due to the difficulties associated with differentiating dead from dormant cells. A number of molecular techniques that may help to distinguish between metabolic states of bacterial cells are presented. The calibration of growth, death and maintenance parameters in rhizosphere models is also discussed. A simple model of rhizosphere carbon flow has been constructed and a sensitivity analysis was carried out on the model to highlight which parameters were most influential when simulating carbon flux. It was observed that the parameters that most heavily influenced long-term carbon compartmentalisation in the rhizosphere were exudation rate and biomass yield. It was concluded that future efforts to simulate carbon flow in the rhizosphere should aim to increase ecological realism in model structure. This revised version was published online in June 2006 with corrections to the Cover Date.     

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。     

植物根际微生物群落构建的研究进展

植物根际是指植物根系与土壤的交界面,是根系自身生命活动和代谢对土壤影响最直接、最强烈的区域,其物理、化学和生物性质不同于土体土壤。在这个区域里,与植物发生相互作用的大量微生物,被称为根际微生物。根际微生物在植物的生长发育和植物病虫害的生物防治等方面都具有十分重要的意义。本文总结了根际微生物群落构建的研究现状,介绍了根际微生物的经典和最新的研究方法,包括根箱法、同位素技术以及高通量测序、菌群定量分析、高通量分离培养等方法在根际微生物研究中的应用,讨论了植物根系分泌物(碳水化物、氨基酸、黄酮类、酚类、激素及其信号物质)和土壤物理化学性质对根际微生物群落的影响,概述了根际微生物-植物的互作机制,以及根际微生物群落对植物的促生作用、提高植物抗逆性和抑制作用,并对根际微生物群落研究中存在的问题和未来发展方向进行了展望。     

连香树人工林根系分泌物输入季节性变化及其驱动的根际微生物特性研究

根系分泌物介导的根际过程具有重要的生态学效应,但目前有关植物尤其是濒危珍稀植物根系分泌物及其介导的土壤生物地球化学循环过程的原位季节动态研究甚少。本文以阿坝州茂县大沟流域30年生的国家二级濒危保护植物连香树人工林为研究对象,于2014年4、7、9和12月利用原位收集装置对连香树根系分泌物进行原位收集;并同步分析了根际土壤微生物特性的变化。结果表明：（1）根系分泌物C和N分泌速率均呈现明显的季节动态变化,其中,夏季最高,冬季最低。（2）根际土壤微生物碳、氮及土壤酶活性显著高于非根际,表现出正的根际效应;根际效应也与根系分泌物输入表现出类似的季节动态规律,即夏季（7月）最高,而冬季（12月）最低。（3）进一步相关分析表明：根系分泌物分泌速率与土壤微生物碳、氮及土壤酶活性根际效应值呈线性正相关,表明根系分泌物输入是驱动根际微生物活性的重要因子,揭示连香树根系碳输入对根际土壤过程和功能的季节响应。未来研究应加强根系分泌物输入与土壤生物地球化学循环过程的偶联效应与机制研究。     

根系分泌物生态学研究

在植物生长过程中 ,由根系的不同部位分泌或溢泌一些无机离子及有机化合物 ,这些物质统称为根系分泌物。植物在其生长过程中不断地分泌无机离子及有机化合物 ,这是植物长期适应外界环境而形成的一种适应机制。早在 2 0世纪 5 0年代就有人对植物根系分泌物进行了研究 ,Ｒｏｖｉｒａ等[4 3] 和Ｖａｎｃｕｒａ等[4 7] 对根土界面根系分泌物进行了系统的研究 ,切尔诺布里维卡[2 6 ] 研究了植物根系分泌物的生物学作用 ,揭示了其在间作中的作用 ,直到 70年代对根系分泌物的研究才出现了蓬勃发展的趋势。近年来的研究表明 ,根系分泌物是保持根际微…     

Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation

The degree to which rising atmospheric CO(2) will be offset by carbon (C) sequestration in forests depends in part on the capacity of trees and soil microbes to make physiological adjustments that can alleviate resource limitation. Here, we show for the first time that mature trees exposed to CO(2) enrichment increase the release of soluble C from roots to soil, and that such increases are coupled to the accelerated turnover of nitrogen (N) pools in the rhizosphere. Over the course of 3 years, we measured in situ rates of root exudation from 420 intact loblolly pine (Pinus taeda L.) roots. Trees fumigated with elevated CO(2) (200 p.p.m.v. over background) increased exudation rates (μg C cm(-1) root h(-1) ) by 55% during the primary growing season, leading to a 50% annual increase in dissolved organic inputs to fumigated forest soils. These increases in root-derived C were positively correlated with microbial release of extracellular enzymes involved in breakdown of organic N (R(2) = 0.66; P = 0.006) in the rhizosphere, indicating that exudation stimulated microbial activity and accelerated the rate of soil organic matter (SOM) turnover. In support of this conclusion, trees exposed to both elevated CO(2) and N fertilization did not increase exudation rates and had reduced enzyme activities in the rhizosphere. Collectively, our results provide field-based empirical support suggesting that sustained growth responses of forests to elevated CO(2) in low fertility soils are maintained by enhanced rates of microbial activity and N cycling fuelled by inputs of root-derived C. To the extent that increases in exudation also stimulate SOM decomposition, such changes may prevent soil C accumulation in forest ecosystems.