植物根系分泌物生态效应及其影响因素研究综述

植物根系分泌物的形成是植物体代谢过程中重要的生理现象,为“植物-土壤”体系物质周转的重要环节.研究植物根系分泌物对于了解陆地生态系统质能过程、碳氮收支平衡及提高生态系统的初级生产具有重要意义.本文从植物根系分泌物对植物生理性状、土壤微生物、土壤物质周转及有机污染物降解影响等4个方面对植物根系分泌物的生态效应进行综述,并从重金属含量、营养元素水平、土壤水分和光热条件、物种基因型、土壤微生物状况和外源有机污染物添加的角度综述了影响植物根系分泌物的因素,旨在对植物根系分泌物的生态效应和影响因素进行总结,并根据目前的研究现状,从研究对象、研究方法和效应评估方面进行了展望.     

植物根系分泌物对土壤污染修复的作用及影响机理

生物修复是一种经济环保的土壤修复技术。根系分泌物是利用生物修复污染土壤过程中的关键物质,也是植物与土壤微生物进行物质交换和信息传递的重要载体,在植物响应污染物胁迫中扮演重要角色。研究植物根系分泌物对土壤污染修复的作用和影响机理,是深入理解植物和微生物环境适应机制的重要途径,对促进生物修复污染土壤有重要指导意义。从污染物胁迫对根系分泌物的影响、根系分泌物对土壤污染物环境行为的影响、根系分泌物在调控污染土壤中根际微生物群落结构和多样性中发挥的作用等几个方面综述了根系分泌物对土壤污染修复的影响及内在机制。研究结果表明,根系分泌物在降低重金属对植物的毒性、加速有机污染物降解等方面有非常重要的作用。根系分泌物对土壤微生物的丰度和多样性均有显著影响,其与根际微生物互作在土壤污染物的消减中发挥了重要的调控作用。在此基础上,提出了以往研究中的不足,并对污染物胁迫下根系分泌物未来研究的方向和趋势进行了展望。     

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。     

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。     

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。     

根系分泌物研究方法(综述)

根系分泌物是植物与土壤进行物质、能量与信息交流的重要载体物质.研究根系分泌物的种类、数量对了解植物与土壤及根际微生物相互作用机理具有重要指导意义,根系分泌物研究方法的选择对研究结果有重要的影响.本文着重讨论根系分泌物收集、分离纯化及鉴定的常用方法,并列举特定根系分泌物的研究方法,旨在为根系分泌物的研究提供借鉴和参考.     

植物根系分泌物在污染及沙化土壤修复中的应用现状与前景

近年来重金属污染等生态环境问题日益受到重视,而物理、化学修复方法存在的诸如成本高、二次污染等问题,使得利用植物、微生物等进行联合治理成为环境修复的重要手段。植物根系分泌物作为植物与土壤进行营养和信息交流的重要媒介,不但对植物的生长具有重要作用,其在污染及沙化土壤修复中作用的研究也得以广泛开展。本文对根系分泌物的组成、分泌机制进行了阐述,并对其在植物吸收重金属、化感作用、植物根系与根际微生物互作、改变土壤理化性质等过程中的作用及机理进行了总结。此外,本文还对利用根系分泌物和根际微生物在生态环境治理中的应用现状、面临的难题及未来的发展等进行了讨论。希望本文可为基于植物与微生物进行的环境修复技术的实际应用提供理论支撑。     

根系分泌物及其在植物修复中的作用

 近年来环境污染日益严重,污染物在土壤植物中的行为引起了人们的高度关注。利用植物去除土壤水体等介质中污染物的植物修复是近10年来兴起的一项安全、廉价的技术,已成为污染生态学和环境生态学的研究热点,它通过植物吸收、根滤、稳定、挥发等方式清除环境中的重金属和有机污染物。国内外有关植物修复的研究报道和概述很多, 但对植物根系分泌物在植物修复中所起的作用及其机理少有述评。 本文从根系分泌物对土壤重金属和土壤有机污染物的去除作用出发,对根系分泌物的种类、数量及其在去除环境污染物中的作用机理和功能地位进行了总结,并借助研究事例对影响植物根系分泌的内外因子,如植物种类、营养胁迫、重金属胁迫、根际环境的理化性质、土壤微生物及其它环境因子进行了讨论。概言之,根系分泌物在修复污染土壤中的重金属途径是多种多样的,主要是通过调节根际pH值、与重金属形成螯合物、络合反应、沉淀、提高土壤微生物数量和活性来改变重金属在根际中的存在形态以及提高重金属的生物有效性,从而减轻它对环境的危害。在清除有机污染物时,根系分泌物中的酶可以对有机污染物进行直接降解,根系分泌物影响下的微生物也可以对有机污染物进行间接降解,且被认为是主要的降解途径。根系分泌物在植物修复过程中确实起着某些重要作用,今后应将这方面的研究重点放在某些特异性根系分泌物植物,尤其是某些重金属超富集植物资源的寻找、筛选上,通过室内实验和野外研究确定其根系分泌物对清除重金属和有机污染物的效率,证实超富集植物根系分泌物的特异性与污染物超富集的内在联系,找到污染土壤生态恢复和治理的有效方法并加以推广应用,如针对性地在被污染地大面积种植此类具特异性根分泌物植物,并辅以营林措施如修剪等,加快生物修复进程,提高修复效率。植物根系分泌物在植物修复过程中所具有的重要生态意义和可能应用前景,为污染生态学和化学生态学之间的联合研究开拓了全新的领域,今后将取得新的突破和重要进展。     

连作障碍与根际微生态研究Ⅱ.根系分泌物与酚酸物质

阐述了作物主要根系分泌物与作物种类、生长期以及与所处环境的关系。并从植物的残体分解、作物根系的分泌等方面论述了土壤中酚酸物质的来源、存在形态、吸附机理及其对作物生长发育与土壤生物活性的影响与机制。     

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。     

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…     

Root exudates and their ecological consequences in forest ecosystems: Problems and perspective

植物根际过程与调控机理研究已成为当前土壤学最活跃、最敏感的研究领域, 而根系分泌物作为根系-土壤-微生物界面物质能量交换和信息传递的重要媒介物质, 是构成根际微生态系统活力与功能特征的内在驱动因素, 是根际概念与根际过程存在的重要前提和基础。然而, 由于传统的根际过程研究更强调以实际生产问题为导向, 加之农作物生长周期较短、操作便利等诸多因素, 以往对植物根系分泌物研究主要聚焦在农业生态系统, 而有关根系分泌物在森林生态系统中的重要作用与调控机理研究甚少, 认识相对零散和片段化。基于此, 该文结合作者实际研究工作中的主要成果和该领域国际前沿动态, 综述了森林根系分泌物的生态重要性, 重点论述了目前森林根系分泌物生态学研究中存在的主要问题与不足, 在此基础上展望了未来森林根系分泌物生态学研究中值得关注的重点方向和研究内容。     

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.     

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     

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     

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     

富钾基因型籽粒苋主要根系分泌物及其对土壤矿物态钾的活化作用

通过试验,研究了2种供K水平对籽粒苋(Amaranthus spp.)富K基因型和一般基因型根系分泌物含量变化的影响,以及在低K胁迫时3个生长期两类基因型主要根系分泌物含量的变化特点,模拟了籽粒苋根系分泌物对土壤矿物态钾的活化作用.结果表明,籽粒苋根系分泌物中可溶性糖、氨基酸和有机酸含量随供K水平的升高而降低,且富K基因型根系分泌物中3种物质的分泌量始终大于一般基因型;在正常供K条件下,两基因型根系分泌能力相近,但在低K处理时,前者显著高于后者,差异显著;在2种供K水平下,根系有机酸分泌量在3种分泌物中占绝对优势,分别是可溶性糖和氨基酸分泌量的几十倍和几百倍.籽粒苋生长到50 d时,一般基因型根系可溶性糖、氨基酸和有机酸的分泌量较40 d时迅速降低.富K基因型根系分泌物中可溶性糖、氨基酸和有机酸含量在3个生长时期均大于一般基因型,且随着生长时间的延长,两基因型间可溶性糖、氨基酸和有机酸含量的差异明显增大.两类基因型在3个生长时期均以分泌有机酸为主,其占总分泌量的93%以上.籽粒苋根系分泌物处理后的土壤速效钾含量均高于清水对照处理,富K基因型在低K胁迫时的根系分泌物对土壤K的活化作用明显大于一般基因型.     

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.     

Evolutionary Divergences in Root Exudate Composition among Ecologically-Contrasting Helianthus Species

Plant roots exude numerous metabolites into the soil that influence nutrient availability. Although root exudate composition is hypothesized to be under selection in low fertility soils, few studies have tested this hypothesis in a phylogenetic framework. In this study, we examined root exudates of three pairs of Helianthus species chosen as phylogenetically-independent contrasts with respect to native soil nutrient availability. Under controlled environmental conditions, seedlings were grown to the three-leaf-pair stage, then transferred to either high or low nutrient treatments. After five days of nutrient treatments, we used gas chromatography-mass spectrometry for analysis of root exudates, and detected 37 metabolites across species. When compared in the high nutrient treatment, species native to low nutrient soils exhibited overall higher exudation than their sister species native to high nutrient soils in all three species pairs, providing support for repeated evolutionary shifts in response to native soil fertility. Species native to low nutrient soils and those native to high nutrient soils responded similarly to low nutrient treatments with increased exudation of organic acids (fumaric, citric, malic acids) and glucose, potentially as a mechanism to enhance nutrition acquisition. However, species native to low nutrient soils also responded to low nutrient treatments with a larger decrease in exudation of amino acids than species native to high nutrient soils in all three species pairs. This indicates that species native to low nutrient soils have evolved a unique sensitivity to changes in nutrient availability for some, but not all, root exudates. Overall, these repeated evolutionary divergences between species native to low nutrient soils and those native to high nutrient soils provide evidence for the adaptive value of root exudation, and its plasticity, in contrasting soil environments.     

Autoinhibition and soil allelochemical (cyclic dipeptide) levels in replanted Chinese fir (Cunninghamia lanceolata) plantations

Aims and background

Despite increasing knowledge of the role of allelochemicals in the productivity decline of replanted Chinese fir plantations, relatively little is known about the levels and sources of allelochemicals in relation to autoinhibition.

Methods

Allelopathic potential of litter, root exudates, and soils in successive rotations of Chinese fir plantations were detected. An allelochemical cyclic dipeptide (6-hydroxy-1,3-dimethyl-8-nonadecyl-[1,4]-diazocane-2,5-dione) from litter, root exudates, and soils in successive rotations was quantified.

Results

Extracts of leaf litter, fine root, and root exudates significantly inhibited the growth of Chinese fir germinants, and inhibition increased with successive rotations. Similar results were observed in the rhizosphere soil, basal soil, and bulk soil. The largest observed inhibition occurred in the rhizosphere soil. Furthermore, cyclic dipeptide was found in litter, root exudates, and soils, and the concentrations increased with successive rotations. The rhizosphere soil had the highest cyclic dipeptide level, followed by basal soil, while bulk soil contained the lowest concentration. There was a significant positive relationship between the inhibition of radicle growth of Chinese fir germinants and the concentration of cyclic dipeptide. Annual release of cyclic dipeptide through root exudation was 2.08–9.78 mol ha^?1 annum, but the annual release of cyclic dipeptide through leaf litter decomposition was lowered to 0.32–1.41 mol ha^?1 annum.

Conclusions

Cyclic dipeptide which caused autoinhibition of Chinese fir may be released into the soil through litter decomposition and root exudation. Root exudates provided more contributions to soil cyclic dipeptide levels than litter in Chinese fir plantations.