How plants communicate using the underground information superhighway

The rhizosphere is a densely populated area in which plant roots must compete with invading root systems of neighboring plants for space, water, and mineral nutrients, and with other soil-borne organisms, including bacteria and fungi. Root-root and root-microbe communications are continuous occurrences in this biologically active soil zone. How do roots manage to simultaneously communicate with neighboring plants, and with symbiotic and pathogenic organisms within this crowded rhizosphere? Increasing evidence suggests that root exudates might initiate and manipulate biological and physical interactions between roots and soil organisms, and thus play an active role in root-root and root-microbe communication.     

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。     

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。     

Root Exudates Regulate Soil Fungal Community Composition and Diversity

Plants are in constant contact with a community of soil biota that contains fungi ranging from pathogenic to symbiotic. A few studies have demonstrated a critical role of chemical communication in establishing highly specialized relationships, but the general role for root exudates in structuring the soil fungal community is poorly described. This study demonstrates that two model plant species (Arabidopsis thaliana and Medicago truncatula) are able to maintain resident soil fungal populations but unable to maintain nonresident soil fungal populations. This is mediated largely through root exudates: the effects of adding in vitro-generated root exudates to the soil fungal community were qualitatively and quantitatively similar to the results observed for plants grown in those same soils. This effect is observed for total fungal biomass, phylotype diversity, and overall community similarity to the starting community. Nonresident plants and root exudates influenced the fungal community by both positively and negatively impacting the relative abundance of individual phylotypes. A net increase in fungal biomass was observed when nonresident root exudates were added to resident plant treatments, suggesting that increases in specific carbon substrates and/or signaling compounds support an increased soil fungal population load. This study establishes root exudates as a mechanism through which a plant is able to regulate soil fungal community composition.     

The root microbiome: Community assembly and its contributions to plant fitness

The root microbiome refers to the community of microbes living in association with a plant's roots, and includes mutualists, pathogens, and commensals. Here we focus on recent advances in the study of root commensal community which is the major research object of microbiome-related researches. With the rapid development of new technologies, plant–commensal interactions can be explored with unprecedented breadth and depth. Both the soil environment and the host plant drive commensal community assembly. The bulk soil is the seed bank of potential commensals, and plants use root exudates and immune responses to build healthy microbial communities from the available microbes. The plant microbiome extends the functional system of plants by participating in a variety of processes, including nutrient absorption, growth promotion, and resistance to biotic and abiotic stresses. Plants and their microbiomes have evolved adaptation strategies over time. However, there is still a huge gap in our understanding of the regulatory mechanisms of plant–commensal interactions. In this review, we summarize recent research on the assembly of root microbial communities and the effects of these communities on plant growth and development, and look at the prospects for promoting sustainable agricultural development through the study of the root microbiome.     

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.     

Polymer and sprinkler droplet energy effects on sugar beet emergence, soil penetration resistance, and aggregate stability

The rapid development of agricultural biotechnology and release of new transgenic plants for agriculture has provided many economic benefits, but has also raised concern over the potential impact of transgenic plants on the environment. Considerable research has now been conducted on the effects of transgenic plants on soil microorganisms. These effects include unintentional changes in the chemical compositions of root exudates, and the direct effects of transgenic proteins on nontarget species of soil microorganisms. Most studies to date suggest that transgenic plants that have been released cause minor changes in microbial community structures that are often transient in duration. However, due to our limited knowledge of the linkage between microbial community structure and function, more work needs to be done on a case-by-case basis to further evaluate the effects of transgenic plants on soil microorganisms and soil ecosystem functions. This review summarizes the results of a variety of experiments that have been conducted to specifically test the effects of transgenic plants on soil microorganisms, and particularly examines the types of methods that are being used to study microbial interactions with transgenic plants.     

植物化学通讯研究进展

 生物的信息传递是生命科学中引人入胜的研究领域之一,生物种间种内和个体内都存在着物理和化学等各种信息交流方式。植物种间种内是否通过物理信号进行通讯交流还是一个未知数,但邻近的同种或异种植物通过化学物质为媒介的通讯关系确是客观存在的。最近,愈来愈多的研究证明：许多陆生植物种可以合成并释放特定的次生物质,这些次生物质可以通过空气和土壤两种载体进行信息传递,尤其是在植物受到侵袭和寄生条件下。茉莉酮酸甲酯、水杨酸甲酯和乙烯等挥发性次生物质被确证为以空气为媒介进行植物种间和种内通讯的化学信号分子。植物根分泌的黄酮和氢醌等分子也可以经土壤媒介传递信息。由于在自然条件下植物根系分泌物的收集和活性信号分子的俘获及鉴定技术还未能突破,这增加了以土壤为媒介的植物种间和种内化学通讯关系研究的难度。但不论如何,植物的化学通讯是植物种间和种内交流的主要方式,植物间的化学通讯关系的研究还处于突破的前夜,这方面的任一研究成果都会引起世界性的关注。因此,破译植物种间和种内化学通讯密码具有重要的学术价值。     

Effects of genetically modified starch metabolism in potato plants on photosynthate fluxes into the rhizosphere and on microbial degraders of root exudates

A high percentage of photosynthetically assimilated carbon is released into soil via root exudates, which are acknowledged as the most important factor for the development of microbial rhizosphere communities. As quality and quantity of root exudates are dependent on plant genotype, the genetic engineering of plants might also influence carbon partitioning within the plant and thus microbial rhizosphere community structure. In this study, the carbon allocation patterns within the plant-rhizosphere system of a genetically modified amylopectin-accumulating potato line (Solanum tuberosum L.) were linked to microbial degraders of root exudates under greenhouse conditions, using (13)C-CO(2) pulse-chase labelling in combination with phospholipid fatty acid (PLFA) analysis. In addition, GM plants were compared with the parental cultivar as well as a second potato cultivar obtained by classical breeding. Rhizosphere samples were obtained during young leaf developmental and flowering stages. (13)C allocation in aboveground plant biomass, water-extractable organic carbon, microbial biomass carbon and PLFA as well as the microbial community structure in the rhizosphere varied significantly between the natural potato cultivars. However, no differences between the GM line and its parental cultivar were observed. Besides the considerable impact of plant cultivar, the plant developmental stage affected carbon partitioning via the plant into the rhizosphere and, subsequently, microbial communities involved in the transformation of root exudates.     

植物与土壤微生物在调控生态系统养分循环中的作用

陆地生态系统的地上、地下是相互联系的。植物与土壤微生物作为陆地生态系统中的重要组成部分, 它们之间的相互作用是生态系统地上、地下结合的重要纽带。该文首先介绍了植物在养分循环中对营养元素的吸收、积累和归还等作用, 阐述了土壤微生物对养分有效性及土壤质量具有重要的作用。其次, 重点综述了植物与土壤微生物之间相互依存、相互竞争的关系。植物通过其凋落物与分泌物为土壤微生物提供营养, 土壤微生物作为分解者提供植物可吸收的营养元素, 比如共生体菌根真菌即可使植物根与土壤真菌达到互惠。然而, 植物的养分吸收与微生物的养分固持同时存在, 因而两者之间存在对养分的竞争。通过植物多样性对土壤微生物多样性的影响分析, 以及土壤微生物直接或间接作用于植物多样性和生产力的分析, 探讨了植物物种多样性与土壤微生物多样性之间的内在联系。针对当前植物与土壤微生物对养分循环的调控机制的争论, 提出植物凋落物是调节植物与土壤微生物养分循环的良好媒介, 植物与土壤微生物的共同作用对维持整个生态系统的稳定性具有重要意义。也指出了目前在陆地生态系统地上、地下研究中存在的不足和亟待解决的问题。     

Mechanisms of microbially enhanced Fe acquisition in red clover (Trifolium pratense L.)

Soil microorganisms may play an important role in plant Fe uptake from soils with low Fe bioavailability, but there is little direct experimental evidence to date. We grew red clover, an Fe-efficient leguminous plant, in a calcareous soil to investigate the role of soil microbial activity in plant Fe uptake. Compared with plants grown in non-sterlie (NS) grown plants, growth and Fe content of the sterile(s) grown plants was significantly inhibited, but was improved by foliar application of Fe EDTA, indicating that soil microbial activity should play an important role in plant Fe acquisition. When soil solution was incubated with phenolic root exudates from Fe-deficient red clover, a few microbial species thrived while growth of the rest was inhibited, suggesting that the Fe-deficient (-Fe) root exudates selectively influenced the rhizosphere's microbial community. Eighty six per cent of the phenolic-tolerant microbes could produce siderophore [the Fe(III) chelator] under -Fe conditions, and 71% could secrete auxin-like compounds. Interestingly, the synthetic and microbial auxins (MAs) significantly enhanced the Ferric reduction system, suggesting that MAs, in addition to siderophores, are important to plant Fe uptake. Finally, plant growth and Fe uptake in sterilized soil were significantly increased by rhizobia inoculation. Root Fe-EDTA reductase activity in the -Fe plant was significantly enhanced by rhizobia infection, and the rhizobia could produce auxin but not siderophore under Fe-limiting conditions, suggesting that the contribution of nodulating rhizobia to plant Fe uptake can be at least partially attributed to stimulation of turbo reductase activity through nodule formation and auxin production in the rhizosphere. Based on these observations, we propose as a model that root exudates from -Fe plants selectively influence the rhizosphere microbial community, and the microbes in turn favour plant Fe acquisition by producing siderophores and auxins.     

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     

Herbivore-induced changes in plant carbon allocation: assessment of below-ground C fluxes using carbon-14

Effects of above-ground herbivory on short-term plant carbon allocation were studied using maize (Zea mays) and a generalist lubber grasshopper (Romalea guttata). We hypothesized that above-ground herbivory stimulates current net carbon assimilate allocation to below-ground components, such as roots, root exudation and root and soil respiration. Maize plants 24 days old were grazed (c. 25–50% leaf area removed) by caging grasshoppers around individual plants and 18 h later pulse-labelled with¹⁴CO₂. During the next 8 h,¹⁴C assimilates were traced to shoots, roots, root plus soil respiration, root exudates, rhizosphere soil, and bulk soil using carbon-14 techniques. Significant positive relationships were observed between herbivory and carbon allocated to roots, root exudates, and root and soil respiration, and a significant negative relationship between herbivory and carbon allocated to shoots. No relationship was observed between herbivory and¹⁴C recovered from soil. While herbivory increased root and soil respiration, the peak time for¹⁴CO₂ evolved as respiration was not altered, thereby suggesting that herbivory only increases the magnitude of respiration, not patterns of translocation through time. Although there was a trend for lower photosynthetic rates of grazed plants than photosynthetic rates of ungrazed plants, no significant differences were observed among grazed and ungrazed plants. We conclude that above-ground herbivory can increase plant carbon fluxes below ground (roots, root exudates, and rhizosphere respiration), thus increasing resources (e.g., root exudates) available to soil organisms, especially microbial populations.     

植物寄生对生态系统结构和功能的影响

寄生植物是生态系统中的特殊类群之一。植物寄生可以驱动生态系统中生物与非生物因子的变化,在生态系统结构与功能中起关键作用。寄生植物可以通过对寄主营养的集聚、改变凋落物的质量与数量、改变根的周转与分泌物格局、改变土壤水势,从而影响土壤理化特性。寄生植物会改变寄主的行为,改变寄主与非寄主植物之间的相互作用,从而影响植物群落的结构、多样性和动态,进而影响植被演替和植被生产力等。寄生植物与寄主均可被消费者取食,可直接或间接地影响生态系统的食草动物,包括草食昆虫等。寄生植物与寄主的其它寄生物存在竞争关系,可以直接或间接地影响寄主的其它寄生植物或病原真菌。寄生植物可以明显地改变土壤地球化学循环,将固有的不可动的成分转变为可利用的营养成分,改变土壤生物群落的结构与功能,从而显著影响地下生物群落。这些表明,植物寄生对生态系统的结构和功能有重要影响。针对特殊的被入侵的植物群落,该地寄生植物可以通过影响入侵植物寄主的生长、繁殖、生物量分配格局,改变土壤的理化特性,促进非寄主的非优势本地植物的生长,从而改变被入侵植物群落结构与多样性,达到生物防治及生态恢复的目的。     

Root exudate cocktails: the link between plant diversity and soil microorganisms?

Higher plant diversity is often associated with higher soil microbial biomass and diversity, which is assumed to be partly due to elevated root exudate diversity. However, there is little experimental evidence that diversity of root exudates shapes soil microbial communities. We tested whether higher root exudate diversity enhances soil microbial biomass and diversity in a plant diversity gradient, thereby negating significant plant diversity effects on soil microbial properties. We set up plant monocultures and two‐ and three‐species mixtures in microcosms using functionally dissimilar plants and soil of a grassland biodiversity experiment in Germany. Artificial exudate cocktails were added by combining the most common sugars, organic acids, and amino acids found in root exudates. We applied four different exudate cocktails: two exudate diversity levels (low‐ and high‐diversity) and two nutrient‐enriched levels (carbon‐ and nitrogen‐enriched), and a control with water only. Soil microorganisms were more carbon‐ than nitrogen‐limited. Cultivation‐independent fingerprinting analysis revealed significantly different soil microbial communities among exudate diversity treatments. Most notably and according to our hypothesis, adding diverse exudate cocktails negated the significant plant diversity effect on soil microbial properties. Our findings provide the first experimental evidence that root exudate diversity is a crucial link between plant diversity and soil microorganisms.     

连作障碍与根际微生态研究 Ⅰ. 根系分泌物及其生态效应

作物、蔬菜、果树以及苗木长期连作后,皆出现生长衰退和产量降低。许多研究结果表明,连作条件下土壤生态环境对植物生长有很大的影响,尤以植物残体与病原微生物的分解产物,对植物有致毒作用,并影响植物根系分泌物正常代谢,以致于发生自毒作用。本文围绕根系分泌物与根际微生态的相互关系,系统地介绍连作障碍条件下,影响根系分泌物的环境因素(土壤空气、湿度、养分与微生物)、活性物质(自身毒素、残体分解物、微生物产生毒素)、土壤病原菌等的根际效应,为深入研究根系分泌物与连作障碍的相互作用机制提供启示。     

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。     

Biotic and Abiotic Properties Mediating Plant Diversity Effects on Soil Microbial Communities in an Experimental Grassland

Plant diversity drives changes in the soil microbial community which may result in alterations in ecosystem functions. However, the governing factors between the composition of soil microbial communities and plant diversity are not well understood. We investigated the impact of plant diversity (plant species richness and functional group richness) and plant functional group identity on soil microbial biomass and soil microbial community structure in experimental grassland ecosystems. Total microbial biomass and community structure were determined by phospholipid fatty acid (PLFA) analysis. The diversity gradient covered 1, 2, 4, 8, 16 and 60 plant species and 1, 2, 3 and 4 plant functional groups (grasses, legumes, small herbs and tall herbs). In May 2007, soil samples were taken from experimental plots and from nearby fields and meadows. Beside soil texture, plant species richness was the main driver of soil microbial biomass. Structural equation modeling revealed that the positive plant diversity effect was mainly mediated by higher leaf area index resulting in higher soil moisture in the top soil layer. The fungal-to-bacterial biomass ratio was positively affected by plant functional group richness and negatively by the presence of legumes. Bacteria were more closely related to abiotic differences caused by plant diversity, while fungi were more affected by plant-derived organic matter inputs. We found diverse plant communities promoted faster transition of soil microbial communities typical for arable land towards grassland communities. Although some mechanisms underlying the plant diversity effect on soil microorganisms could be identified, future studies have to determine plant traits shaping soil microbial community structure. We suspect differences in root traits among different plant communities, such as root turnover rates and chemical composition of root exudates, to structure soil microbial communities.     

Molecular dialogues between Trichoderma and roots: Role of the fungal secretome

Trichoderma species are opportunistic fungi residing primarily in soil, tree bark and on wild mushrooms. Trichoderma is capable of killing other fungi and penetrating plant roots, and is commonly used as both a biofungicide and inducer of plant defence against pathogens. These fungi also exert other beneficial effects on plants including growth promotion and tolerance to abiotic stresses, primarily mediated by their intimate interactions with roots. In root–microbe interactions (both beneficial and harmful), fungal secreted proteins play a crucial role in establishing contact with the roots, fungal attachment, root penetration and triggering of plant responses. In Trichoderma–root interactions, the sucrose present in root exudates has been demonstrated to be important in fungal attraction. Attachment to roots is mediated by hydrophobin-like proteins, and secreted swollenins and plant cell wall degrading enzymes facilitate internalization of the fungal hyphae. During the early stage of penetration, suppression of plant defence is vital to successful initial root colonisation; this is mediated by small soluble cysteine-rich secreted proteins (effector-like proteins). Up to this stage, Trichoderma's behaviour is similar to that of a plant pathogen invading root structures. However, subsequent events like oxidative bursts, the synthesis of salicylic acid by the plants, and secretion of elicitor-like proteins by Trichoderma spp. differentiate this fungus from pathogens. These processes induce immunity in plants that help counter subsequent invasion by plant pathogens and insects. In this review, we present an inventory of soluble secreted proteins from Trichoderma that might play an active role in beneficial Trichoderma–plant interactions, and review the function of such proteins where known.