Soil microorganisms interacting with residue-derived allelochemicals effects on seed germination

Despite the knowledge regarding allelopathy, known as a major ecological mechanism for biological weed control, had increased greatly, the role of soil microorganisms in that field remained controversial. The study sought to evaluate the interference potential of soil microorganisms, residues-derived allelochemicals and their interaction on seed germination and understand the variation of microbial community in allelopathic activities. Three different rice residues-derived fractions from variety PI312777 (extracts, straw fraction and fresh residue) were applied to sterile and live soils to disentangle the interference potential of soil microorganisms, residues-derived allelochemicals and their interaction concerned allelopathic activities. The results demonstrated that microbe-only and residues-only exerted onefold promotion and inhibition effects on lettuce (Lactuca sativa Linn.) seed germination, respectively, whereas, microbe-by-residues interaction showed an inhibition at the beginning, and a feeble promotion later. The 20 most dominant genera of microbes were classified into three clusters, with 13 genera in one cluster, only 1 in the second cluster and 6 in the third one. The genera in the first cluster commonly exerted negative effects on phenol content, while showed positive correlation with seed germination. Interestingly, Bacillus, clustered in the second cluster, had an opposite effect alone. The third cluster genera somehow had a weak correlation with both germination as well as the release of the allelochemicals. Overall, we incorporated molecular methodology for tracking bacterial impacts during incubation with allelochemicals, and demonstrated the mutable role of soil microbes in allelopathy. It may be potentially important for stimulating the beneficial roles of microbes for environmentally friendly weed management.     

Unravelling the beneficial role of microbial contributors in reducing the allelopathic effects of weeds

The field of allelopathy is one of the most fascinating but controversial processes in plant ecology that offers an exciting, interdisciplinary, complex, and challenging study. In spite of the established role of soil microbes in plant health, their role has also been consolidated in studies of allelopathy. Moreover, allelopathy can be better understood by incorporating soil microbial ecology that determines the relevance of allelopathy phenomenon. Therefore, while discussing the role of allelochemicals in plant–plant interactions, the dynamic nature of soil microbes should not be overlooked. The occurrence and toxicity of allelochemicals in soil depend on various factors, but the type of microflora in the surroundings plays a crucial role because it can interfere with its allelopathic nature. Such microbes could be of prime importance for biological control management of weeds reducing the cost and ill effects of chemical herbicides. Among microbes, our main focus is on bacteria—as they are dominant among other microbes and are being used for enhancing crop production for decades—and fungi. Hence, to refer to both bacteria and fungi, we have used the term microbes. This review discusses the beneficial role of microbes in reducing the allelopathic effects of weeds. The review is mainly focused on various functions of bacteria in (1) reducing allelopathic inhibition caused by weeds to reduce crop yield loss, (2) building inherent defense capacity in plants against allelopathic weed, and (3) deciphering beneficial rhizospheric process such as chemotaxis/biofilm, degradation of toxic allelochemicals, and induced gene expression.     

植物化感物质及化感潜力与土壤养分的相互影响

植物化感作用与许多生态因子有关.土壤养分缺乏,影响着许多植物化感物质的产生,从而影响植物的化感潜力;反过来,植物化感物质也通过络合、吸附、酸溶解、竞争、抑制等方式影响土壤的养分形态和水平.本文总结了植物化感物质及化感潜力与土壤养分的相互影响,并提出了今后该领域值得进一步研究的问题.包括以下几方面：加强植物化感研究与土壤 植物营养学研究的结合,以更深入地阐明植物化感物质、化感作用与土壤养分变化的关系;加强植物化感研究与生态系统养分循环研究的结合,以类似自然（nature-like）的方式模拟自然界植物所受的养分干,使养分干扰的化感研究结果更加逼真、可靠;加强对养分过量及受污染时植物化感作用的研究,为揭示农业和林业生产中植物的相互作用机制和生物量变化机制提供新的思路,为生态保护提供科学依据.     

Context dependency of the allelopathic effects of Lonicera maackii on seed germination

Allelopathic effects of invasive plants on native flora may be mitigated by the abiotic and biotic environment into which the allelochemicals are released. Lonicera maackii (Amur honeysuckle), an invasive plant of the eastern deciduous forest, suppresses seed germination in laboratory assays. We investigated how L. maackii leachate interacts with abiotic conditions and with the soil microbial community. First, we tested the effects of leaf extract from L. maackii on germination of the native woodland herb, Blephilia hirsuta, under different light and soil conditions. We found that germination of Blephilia hirsuta was reduced by L. maackii extract, but abiotic conditions did not interact with this effect. We also tested the effects of leaf extract on germination of five native woodland species and L. maackii placed in sterile or live soil. There was an overall suppressive effect of L. maackii extract on itself and the other five native species tested. However, L. maackii extract interacted with live soil in ways that differed with the species being tested and, in some cases, changed over time. Our results indicate that allelopathic potential of L. maackii shows context dependency with respect to soil microorganisms and native species identity but not to light conditions or soil type. Our results imply that restoration of invaded areas may require active reintroduction of species sensitive to allelopathy in live soil. Further, laboratory assays of allelopathy should consider the interaction of allelochemicals with biotic and abiotic conditions to more accurately predict the impacts of allelopathy on plant communities.     

The Impact of Microbial Biotransformation of Catechin in Enhancing the Allelopathic Effects of Rhododendron formosanum

Rhododendron formosanum is distributed widely in the central mountains in Taiwan and the major allelopathic compound in the leaves has been identified as (-)-catechin, which is also a major allelochemical of an invasive spotted knapweed in North America. Soil microorganisms play key roles in ecosystems and influence various important processes, including allelopathy. However, no microorganism has been identified as an allelochemical mediator. This study focused on the role of microorganisms in the allelopathic effects of R. formosanum. The microorganism population in the rhizosphere of R. formosanum was investigated and genetic analysis revealed that the predominant genera of microorganisms in the rhizosphere of R. formosanum were Pseudomonas, Herbaspirillum, and Burkholderia. The dominant genera Pseudomonas utilized (-)-catechin as the carbon source and catalyzed the conversion of (-)-catechin into protocatechuic acid in vitro. The concentrations of allelochemicals in the soil were quantified by liquid chromatography-electrospray ionization/tandem mass spectrometry. The concentration of (-)-catechin in the soil increased significantly during the extreme rainfall in the summer season and suppressed total bacterial populations. Protocatechuic acid accumulation was observed while total bacterial populations increased abundantly in both laboratory and field studies. Allelopathic interactions were tested by evaluating the effects of different allelochemicals on the seed germination, radicle growth, and photosynthesis system II of lettuce. Protocatechuic acid exhibited higher phytotoxicity than (-)-catechin did and the effect of (-)-catechin on the inhibition of seed germination was enhanced by combining it with protocatechuic acid at a low concentration. This study revealed the significance of the allelopathic interactions between R. formosanum and microorganisms in the rhizosphere. These findings demonstrate that knowledge regarding the precise biotransformation process of (-)-catechin by microorganisms in the environment is necessary to increase our understanding of allelopathy.     

Microbial Community Changes Elicited by Exposure to Cyanobacterial Allelochemicals

An increasing body of evidence points out that allelopathy may be an important process shaping microbial communities in aquatic ecosystems. Cyanobacteria have well-documented allelopathic properties, mainly derived from the evaluation of the activity of allelopathic extracts or pure compounds towards monocultures of selected target microorganisms. Consequently, little is known regarding the community dynamics of microorganisms associated with allelopathic interactions. In this laboratory-based study, a Microcystis spp.-dominated microbial community from a freshwater lake was exposed, for 15 days, to exudates from the cyanobacterium Oscillatoria sp. strain LEGE 05292 in laboratory conditions. This cyanobacterium is known to produce the allelochemicals portoamides, which were among the exuded compounds. The community composition was followed (by means of polymerase chain reaction followed by denaturing gradient gel electrophoresis and microscopic analyses) and compared to that of a non-exposed situation. Following exposure, clear differences in the community structure were observed, in particular for cyanobacteria and unicellular eukaryotic taxa. Interestingly, distinct Microcystis genotypes present in the community were differentially impacted by the exposure, highlighting the fine-scale dynamics elicited by the exudates. These results support a role for cyanobacterial allelochemicals in the structuring of aquatic microbial communities.     

小麦化感作用研究进展

小麦是世界第一大粮食作物,在农业生产中占有重要地位．然而,由于人们为保证小麦产量往往施用大量的除草剂和杀菌剂,对环境造成了极大的危害．小麦化感作用是利用小麦活体或残体向环境中释放次生代谢物质对自身或其他生物产生作用,它克服了除草剂和杀菌剂等引起的环境污染问题,具有抑制杂草控制病害的潜力．本文对已有的小麦化感作用的研究进展情况进行了综合评述．其中小麦对杂草、虫害及病害产生防御功能的主要化感物质为异羟肟酸和酚酸类物质．小麦化感物质活性的发挥除了取决于化感物质的种类外,还由小麦自身的遗传因素、环境因素和生物因素的共同作用所决定．小麦化感物质在根际土壤中的滞留、迁移和转化过程、小麦化感作用与土壤生物的关系以及相关的作用机理是小麦化感作用研究的薄弱环节。其研究方法还需进一步探索改进．小麦化感作用在植物保护、环境保护以及作物育种等方面具有广泛的应用前景,促进了小麦抗逆性的增强以及产量和品质的提高．     

Allelopathy in Northern Temperate and Boreal Semi-Natural Woodland

Allelopathy has been widely investigated in forest ecosystems around the world. This ecological process takes part in the vegetation dynamics and partially explains the success or failure of seedling establishment and survival. Therefore, allelopathy could be a common cause of natural regeneration failure, due to adverse impact of understory species directly on trees species (inhibition of germination and seedlings primary growth). This problem has been investigated mainly in coniferous forests, but it also occurs in deciduous forests. Furthermore, allelopathy can involve forest soil microorganisms. They can act as allelochemical mediators, where they are able to inactivate these compounds by means of metabolization, as they are able to produce biologically active compounds using inactive molecules as a substrate. Special attention is given to mycorrhizal fungi and mycor-rhizae, because, in theory, they are able to protect seedlings against allelopathy. Applied aspects of allelopathy in forest ecosystems are also presented, with two main goals: (1) how to reduce the adverse impact of such interactions (chemical and mechanical control of allelochemicals plant producer, afforestation with selected mycorrhiza), but also (2) how to use allelopathy as a tool for forest management. Suggestions to improve the study of allelopathy in forest ecosystems and prospects for future investigations conclude this review.     

小麦化感作用研究进展

小麦是世界第一大粮食作物,在农业生产中占有重要地位.然而,由于人们为保证小麦产量往往施用大量的除草剂和杀菌剂,对环境造成了极大的危害.小麦化感作用是利用小麦活体或残体向环境中释放次生代谢物质对自身或其他生物产生作用,它克服了除草剂和杀菌剂等引起的环境污染问题,具有抑制杂草控制病害的潜力.本文对已有的小麦化感作用的研究进展情况进行了综合评述.其中小麦对杂草、虫害及病害产生防御功能的主要化感物质为异羟肟酸和酚酸类物质.小麦化感物质活性的发挥除了取决于化感物质的种类外,还由小麦自身的遗传因素、环境因素和生物因素的共同作用所决定.小麦化感物质在根际土壤中的滞留、迁移和转化过程、小麦化感作用与土壤生物的关系以及相关的作用机理是小麦化感作用研究的薄弱环节,其研究方法还需进一步探索改进.小麦化感作用在植物保护、环境保护以及作物育种等方面具有广泛的应用前景,促进了小麦抗逆性的增强以及产量和品质的提高.     

Ecophysiological Approach in Allelopathy

Allelopathy is a process that can be present in many ecosystems, according to the literature. Nevertheless, the authors think that, due to the evolutionary constraints, especially from a co-evolution point of view, this process cannot be very extended except in some conditions. Allelopathy can be important when an invader plant affects to the autochthonous species, when soil microorganisms cannot cope with a new molecule and target plants have not co-evolved. It can be important in some conditions when there is a continuous release of allelochemicals into the environment or when there is a very limited metabolism of those substances. Most agricultural and forestry practices tend to increase the possibilities of such a process to be important. Allelopathy can be effective only when plants are in stress due to other mechanisms, for example, when there is a lack of water or competition for nutrients or light is strong. In those conditions, allelochemicals production has been shown to increase. Sometimes there are allelochemicals that are not normally produced if the plant is not under stress. Under stress, the target plant is also more susceptible to the effect of the released phytotoxins. A multidisciplinary ecophysiological approach is needed in studying allelopathy in conjunction with other mechanisms that affect plants. The study can range from the ecological level to the molecular one. New methods are requested to separate the effects of allelopathy from competition.     

Challenges,achievements and opportunities in allelopathy research

Abstract

Allelopathy is defined as the suppression of any aspect of growth and/or development of one plant by another through the release of chemical compounds. Although allelopathic interference has been demonstrated many times using in vitro experiments, few studies have clearly demonstrated allelopathy in natural settings. This difficulty reflects the complexity in examining and demonstrating allelopathic interactions under field conditions. In this paper we address a number of issues related to the complexity of allelopathic interference in higher plants: These are: (i) is a demonstrated pattern or zone of inhibition important in documenting allelopathy? (ii) is it ecologically relevant to explain the allelopathic potential of a species based on a single bioactive chemical? (iii) what is the significance of the various modes of allelochemical release from the plant into the environment? (iv) do soil characteristics clearly influence allelopathic activity? (v) is it necessary to exclude other plant interference mechanisms?, and (vi) how can new achievements in allelopathy research aid in solving problems related to relevant ecological issues encountered in research conducted upon natural systems and agroecosystems? A greater knowledge of plant interactions in ecologically relevant environments, as well as the study of biochemical pathways, will enhance our understanding of the role of allelopathy in agricultural and natural settings. In addition, novel findings related to the relevant enzymes and genes involved in production of putative allelochemicals, allelochemical persistence in the rhizosphere, the molecular target sites of allelochemicals in sensitive plant species and the influence of allelochemicals upon other organisms will likely lead to enhanced utilization of natural products for pest management or as pharmaceuticals and nutraceuticals. This review will address these recent findings, as well as the major challenges which continue to influence the outcomes of allelopathy research.     

Plant phenolics in allelopathy

Phenolics are one of the many secondary metabolites implicated in allelopathy. To establish that allelopathy functions in a natural ecosystem, the allelopathic bioassay must be ecologically realistic so that responses of appropriate bioassay species are determined at relevant concentrations. It is important to isolate, identify, and characterize phenolic compounds from the soil. However, since it is essentially impossible to simulate exact field conditions, experiments must be designed with conditions resembling those found in natural systems. It is argued that allelopathic potential of phenolics can be appreciated only when we have a good understanding of 1) species responses to phenolic allelochemicals, 2) methods for extraction and isolation of phenolic allelochemicals, and 3) how abiotic and biotic factors affect phenolic toxicity.     

Plant allelochemical interference or soil chemical ecology?

While allelopathy has been defined as plant-plant chemical interference, there has been much confusion about what the concept encompasses and how important it is in nature. We distinguish between (1) direct plant-plant interference mediated by allelochemicals, and (2) the effects of secondary compounds released by plants on abiotic and biotic soil processes that affect other plants.It very difficult to demonstrate direct effects of chemicals released by a plant on nearby plants. Although soil ecology-mediated effects of secondary plant compounds do not fit the classical concept of allelopathy, we find support in the literature for the hypothesis that the most important effects of compounds released into the soil environment by plants on other plants occur through such indirect effects. The emphasis on, and skepticism of, direct plant-plant allelopathic interference has led some researchers to demand unreasonably high standards of evidence for establishing even the existence of allelopathic interactions, standards that are not demanded for other plant-plant interactions such as resource competition. While the complete elucidation of the mechanisms by which allelochemicals function in the field is many years away, such elucidation is not necessary to establish the existence of allelopathic interactions.We propose that most of the phenomena broadly referred to as allelopathic interference are better conceptualized and investigated in terms of soil chemical ecology. Even when direct plant-plant allelochemical interference occur, the levels of allelochemicals in the environment and their effects on plants are heavily influenced by abiotic and biotic components of the soil ecosystem. Putting allelopathy in the context of soil ecology can further research and reduce some of the less fruitful controversy surrounding the phenomenon.     

水稻化感作用及其分子生态学研究进展

综述了近年来国际上研究水稻化感作用的新进展,比较分析了当前常用于室内评价水稻化感作用潜力的几种生物测试法的优缺点,指出了琼脂迟播共培法是较为理想的室内生物测试法并已广泛应用于化感作用研究中。在此基础上,分析了水稻化感作用的数量遗传特性及其QTL定位的研究现状;阐明了水稻化感作用的遗传多样性及其分子生态特性;并就当前普遍关注的焦点问题：逆境条件（如低氮或高伴生杂草密度胁迫）常引起水稻化感作用潜力增大的生理过程与分子机制作了阐述。结合近年来应用差异蛋白组学和生物信息学的研究实例,阐明了逆境引起水稻化感作用增强与其酚类合成代谢相关酶蛋白表达丰度增加,萜类合成代谢相关酶蛋白表达丰度下降有关。就究竟什么是水稻的化感物质及其作用方式等问题作了分析与讨论．指出水稻的化感抑草效应是其众多化感物质综合作用的结果,应重视区分化感物质对靶标杂草的原生作用和进入土壤生态系统中经生物转化后的次生作用。根据当前植物化感作用研究的发展趋势,阐明了进一步研究水稻化感作用的焦点问题,提出了水稻化感作用的根际生物学特性与分子生态学机制,是未来国际上竞相角逐的重点研究领域,并认为以现代系统生物学理论为指导,运用基因组学、蛋白质组学和代谢组学等技术方法,是揭示这一分子生态学过程与机制的重要技术选择和优先研究领域。     

Perspectives on allelopathic disruption of plant mutualisms: a framework for individual- and population-level fitness consequences

Mutualisms with mycorrhizal fungi, pollinators, and seed dispersers are critical for plant survival and reproduction. However, mutualism effectiveness is highly sensitive to disturbance by environmental stressors. Allelopathy is often overlooked, yet likely important, as a potential stress on plant mutualism function. Allelochemicals can affect plant mutualisms by either directly interfering with the plant’s ability to produce resources and rewards for its mutualistic partners or by directly or indirectly altering the non-plant mutualist’s behavior. Here we explore the potential effects of allelochemicals on plant mutualisms. Since allelochemicals can reduce plant growth and carbon acquisition, we suggest that allelopathy could directly diminish: (1) carbon provisioning to mycorrhizal fungi, (2) flower, pollen, and nectar production for pollinators, and (3) fruit attractiveness to seed dispersers. Similarly, allelochemicals that directly affect mycorrhizal fungi functioning can reduce the flow of soil resources to their plant partner. Further, volatile allelochemicals or uptake of allelochemicals from the soil by the plant could alter pollen/nectar or fruit attractiveness and indirectly influence pollinator and seed disperser behavior. Finally, we explore the extent to which plant-produced chemicals could have a direct or indirect positive effect on plant mutualisms. We end using these questions to frame future avenues of research that could help to move studies of allelopathy into the broader ecological context of mutualisms.     

The fungal fast lane: common mycorrhizal networks extend bioactive zones of allelochemicals in soils

Allelopathy, a phenomenon where compounds produced by one plant limit the growth of surrounding plants, is a controversially discussed factor in plant-plant interactions with great significance for plant community structure. Common mycorrhizal networks (CMNs) form belowground networks that interconnect multiple plant species; yet these networks are typically ignored in studies of allelopathy. We tested the hypothesis that CMNs facilitate transport of allelochemicals from supplier to target plants, thereby affecting allelopathic interactions. We analyzed accumulation of a model allelopathic substance, the herbicide imazamox, and two allelopathic thiophenes released from Tagetes tenuifolia roots, by diffusion through soil and CMNs. We also conducted bioassays to determine how the accumulated substances affected plant growth. All compounds accumulated to greater levels in target soils with CMNs as opposed to soils without CMNs. This increased accumulation was associated with reduced growth of target plants in soils with CMNs. Our results show that CMNs support transfer of allelochemicals from supplier to target plants and thus lead to allelochemical accumulation at levels that could not be reached by diffusion through soil alone. We conclude that CMNs expand the bioactive zones of allelochemicals in natural environments, with significant implications for interspecies chemical interactions in plant communities.     

Roles of Allelopathy in Plant Biodiversity and Sustainable Agriculture

Allelopathic compounds are metabolites released from plants that might be beneficial or detrimental to the growth of receptor plants. These compounds are involved in the environmental complex of managed or natural ecosystems. Allelopathic compounds have been shown to play important roles in the determination of plant diversity, dominance, succession, and climax of natural vegetation and in the plant productivity of agroecosystems. The overuse of synthetic agrochemicals often causes environmental hazards, an imbalance of soil microorganisms, nutrient deficiency, and change of soil physicochemical properties, resulting in a decrease of crop productivity. The incorporation of allelopathic substances into agricultural management may reduce the use of synthetic herbicides, fungicides, and insecticides and lessen environmental deterioration. Scientists in many different habitats around the world have demonstrated the above examples previously. It is known that most volatile compounds, such as terpenoids, are released from plants in drought areas. In contrast, water-borne phytotoxins, such as phenolics, flavonoids, or alkaloids, are released from plants in humid zone areas. Both allelopathy and autointoxication play an important mechanism in regulating plant biodiversity and plant productivity. A unique case study of a pasture-forest intercropping system, which is particularly emphasized here, could be used as a model for forest management. After the deforestation of coniferous or hardwood forests, a pasture grass, kikuyu grass (Pennisetum clandestinum), was transplanted onto the land. The grass was quickly established within 6 months. Significant suppression of weed growth by the kikuyu grass was found; however, the growth of coniferous or hardwood plants was not suppressed but stimulated. This example as well as others described in this text clearly indicate that allelopathy plays a significant role in sustainable agriculture. Nevertheless, room for allelopathic research in the next century is available for biologists, biochemists, biotechnologists, and chemists. Future allelopathic research should focus on the following tasks: (1) a continuous survey of potential allelochemicals from natural vegetation or microorganisms, (2) the establishment of practical ways of using allelochemicals in the field, (3) to understand the mode of action of allelopathic chemicals in receptor organisms, (4) to understand the role of allelopathic chemicals in biodiversity and ecosystem function, (5) to explore advanced biotechnology for allocating allelopathic chemical genes in plants or microorganisms for biological control, and (6) to challenge the natural product chemists to develop a better methodology for isolating allelopathic compounds or their degraded compounds from the environment, particularly the soil environment.     

植物化感物质活性变化及其作用机理研究进展

本文综述了化感物质活性的变化、变化的原因及其功能机理.化感物质活性在自然过程中,由于温度、光周期、水和土壤等的不同而变化,在功能过程中随其初始浓度、化合物结构和混合程度不同而变化,在植物体内由于组织器官和成熟程度不同而变化,而研究技术和操作过程也影响化感物质的活性.文章还讨论了未来化感作用研究的发展方向.未来化感作用研究将集焦于如下五方面:(1)更有效地在实践中,特别是在农业生产过程中鉴定和提纯化感物质;(2)化感作用在分子结构水平的功能定位;(3)应用化感作用解释植物种间相互作用;(4)化感作用在植被演替过程中的驱动力作用;(5)化感作用在进化过程中的意义.     

On laboratory bioassays in allelopathy

Allelopathy involves the complex chain of chemical communications among plants, including microbes. Laboratory bioassays constitute a significant part of allelopathic research, and various bioassays have been proposed to demonstrate allelopathy under controlled lab conditions. However, many lab bioassays have little or no correspondence to field interaction, which may be due to dissimilarity of the conditions of lab bioassay to natural conditions, lack of standardized techniques, or absence of critical controls. Here we discuss several lab bioassays presently used in allelopathic research for their suitability to demonstrate allelopathy of ecological relevance. We recommend avoiding certain practices, such as grinding plant material to evaluate allelopathic potential and isolation of allelochemicals, using seed germination as the only criterion of growth response, using sand, agar, or autoclaved soil, using organic solvents as extractants in allelopathic bioassays, and eliminating microbial involvement. Care should be taken in the lab to simulate natural conditions and attention should be given to habit, habitat, and life cycle pattern of the allelopathic plants during designing of lab bioassays.     

浮游植物的化感作用

生物化感作用研究是近年来兴起的交叉学科,是化学生态学研究的重要领域。研究水域浮游植物化感作用对了解浮游植物之间、浮游植物与其他生物之间的相互作用及作用机理具有重要意义,对了解赤潮和水华的发生机制及其生态控制等具有非常重要的作用。综述了海洋和湖泊浮游植物化感作用和化感物质的内涵,讨论了水体浮游植物化感作用的特点、研究化感作用的基本方法、化感物质的种类以及影响化感物质作用的生物和非生物因素,详细介绍了浮游植物化感物质的作用机理以及逃避和拈抗化感作用的方式,同时对目前研究的热点问题及未来研究的方向做了简要概述。