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.     

Soil Microorganisms: An Important Determinant of Allelopathic Activity

Current evidence illustrates the significance of soil microbes in influencing the bioavailability of allelochemicals. This review discusses (i) the significance of soil microorganisms in influencing allelopathic expression, (ii) different ways of avoiding microbial degradation of putative allelochemicals, and (iii) the need of incorporating experiments on microbial modification of allelochemicals in laboratory bioassays for allelopathy. Several climatic and edaphic factors affect the soil microflora; therefore, allelopathy should be assessed in a range of soil types. Allelopathy can be better understood in terms of soil microbial ecology, and appropriate methodologies are needed to evaluate the roles of soil microorganisms in chemically-mediated interactions between plants.     

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.     

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.     

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.     

菊科植物化感作用研究进展

对菊科植物化感作用的研究进展进行了综述。菊科植物中至少有 39个属存在化感作用 ,特别是一枝黄花属、向日葵属、胜红蓟属、银胶菊属、蒿属植物等有较多的研究报道。鉴定出的化感物质多为萜类、聚乙炔类、酚类、有机酸类等 ,这些化感物质对多种受体植物表现出程度不同的抑制或促进的效应。其化感作用机理表现在破坏受体膜系统的稳定性及水分平衡关系、抑制氧化磷酸化、促进或阻滞叶绿素的合成、影响矿质元素的吸收利用等。并对菊科植物化感物质在植物生长调节剂、天然除草剂和生物杀虫剂 ,或人工合成除草剂和杀虫剂上应用的前景进行了探讨。本文显示菊科植物的化感作用将在控制外来恶性杂草及维护生态平衡上扮演重要的角色。在当前菊科植物化感作用研究的基础上 ,提出了进一步研究的 6个方向 :(1)化感物质的生物合成途径与关键酶的特性研究 ;(2 )具化感潜势物种资源的调查评价及利用研究 ;(3)化感作用在自然生态系统中的演变规律 ;(4 )菊科重要作物自毒的生化机制及克服途径 ;(5 )具应用前景的菊科植物化感关键酶的基因克隆和转基因 ,并对受体植物基因的表达与调控进行研究 ;(6 )化感作用在可持续发展农业应用上的研究与开发 ,特别是作为天然除草剂及杀虫剂     

植物化感物质对种子萌发的影响

所有的化感物质都对一些植物的种子萌发产生影响.本文介绍了植物化感作用和化感物质的定义,化感物质的释放方式和种类,然后重点综述了化感物质影响种子萌发的作用机理、内外因素和生态学意义.化感物质通过影响细胞膜透性、细胞分裂生长和分化、呼吸作用、蛋白质合成、基因表达、激素合成和平衡,最终影响种子萌发.化感物质对种子萌发的影响与化感物质的种类、浓度、受体植物种类和环境条件等有关.     

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．     

侧柏不同器官水提取物对油松种子萌发和幼苗生长的他感效应

研究了侧柏（Platycladus orientalis）根、枝、叶、果实不同浓度水提取物及枝叶挥发油对油松种子萌发和幼苗生长的他感效应。结果表明：侧柏各器官及挥发油都含有化感物质,均对油松种子萌发及幼苗生长表现出“高抑低促”效应,而且低浓度时的促进作用明显强于高浓度时的抑制作用。与对照相比,在浓度为2 mg·mL^-1时,侧柏根水提取物对油松幼苗根长和鲜重的促进能力最强,其根长和鲜重分别比对照提高了79.78%和376.60%;侧柏叶水提取物对油松幼苗苗高促进能力最强,比对照提高了102.41%;侧柏枝水提取物对油松种子发芽的促进能力最强,发芽率比对照提高了65.63%。而在浓度为30 mg·mL^-1时,除了侧柏根和枝水提取物对油松种子萌发（87.50%）和鲜重（85.11%）表现出明显的抑制作用外,其它提取物对油松幼苗根长和苗高生长的抑制作用均不明显。侧柏枝叶挥发油对油松种子萌发,幼苗生长均表现为促进作用,特别对油松种子萌发具有显著的促进作用,其促进率达70.37%。说明侧柏和油松混交,可能有利于油松树木生长及生产力的提高。     

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.     

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.     

Allelopathy in wheat (Triticum aestivum)

Wheat (Triticum aestivum) allelopathy has potential for the management of weeds, pests and diseases. Both wheat residue allelopathy and wheat seedling allelopathy can be exploited for managing weeds, including resistant biotypes. Wheat varieties differ in allelopathic potential against weeds, indicating that selection of allelopathic varieties might be a useful strategy in integrated weed management. Several categories of allelochemicals for wheat allelopathy have been identified, namely, phenolic acids, hydroxamic acids and short‐chain fatty acids. Wheat allelopathic activity is genetically controlled and a multigenic model has been proposed. Research is underway to identify genetic markers associated with wheat allelopathy. Once allelopathic genes have been located, a breeding programme could be initiated to transfer the genes into modern varieties for weed suppression. The negative impacts of wheat autotoxicity on agricultural production systems have also been identified when wheat straws are retained on the soil surface for conservation farming purposes. A management package to avoid such deleterious effects is discussed. Wheat allelopathy requires further study in order to maximise its allelopathic potential for the control of weeds, pests and diseases, and to minimise its detrimental effects on the growth of wheat and other crops.     

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。     

黄瓜种子及其萌发期的化感作用研究

以大白菜、萝卜、番茄和黄瓜种子为受体,采用实验室培养皿种子发芽生物测试法研究了黄瓜种子浸提液、种子萌发、胚根和芽苗分泌物、芽苗腐解物和芽苗浸提液的化感效应。结果表明:(1)黄瓜种子浸提液对大白菜、萝卜、番茄和黄瓜种子萌发均有化感抑制作用,即黄瓜种子内含有某些化感抑制物质。(2)在水浸提过的黄瓜种子萌发过程中,它不仅对其近邻套种的大白菜、萝卜和番茄种子萌发产生化感抑制作用,而且其胚根和芽苗分泌物对后茬播种的4种蔬菜种子发芽也表现出不同程度的化感抑制作用;黄瓜芽苗腐解物和芽苗水浸提液也对各受体蔬菜种子发芽与生长产生不同程度的化感抑制作用,且随着腐解芽苗量的增加或浸提液浓度的升高,各受体蔬菜种子的发芽指标值、化感效应指数值和综合效应值随之降低。(3)黄瓜种子浸提液及芽苗各器官的化感物质对黄瓜种子的萌发与生长产生了自毒作用,且黄瓜芽苗腐解物、芽苗浸提液、胚根及芽苗分泌物对受体黄瓜的自毒作用均为最大。研究发现,黄瓜种子浸提液、种子萌发时期以及芽苗各器官的化感物质主要是通过抑制受体胚根的生长而起化感抑制作用,即受体蔬菜种子胚根对化感效应最为敏感;因黄瓜种子及萌发期释放化感物质的途径有所不同,导致受体大白菜、萝卜、黄瓜和番茄的化感响应也不相同;在黄瓜种子萌发和芽苗生长的早期,化感物质即开始在芽苗体内进行合成与积累,一部分可通过胚根和芽苗分泌途径释放到环境中,另一部分可通过芽苗腐解途径释放化感物质,并对受体蔬菜种子萌发与生长表现出较强的化感抑制作用。     

Is Allelopathic Activity of Ipomoea murucoides Induced by Xylophage Damage?

Herbivory activates the synthesis of allelochemicals that can mediate plant-plant interactions. There is an inverse relationship between the activity of xylophages and the abundance of epiphytes on Ipomoea murucoides. Xylophagy may modify the branch chemical constitution, which also affects the liberation of allelochemicals with defense and allelopathic properties. We evaluated the bark chemical content and the effect of extracts from branches subjected to treatments of exclusion, mechanical damage and the presence/absence of epiphytes, on the seed germination of the epiphyte Tillandsia recurvata. Principal component analysis showed that branches without any treatment separate from branches subjected to treatments; damaged and excluded branches had similar chemical content but we found no evidence to relate intentional damage with allelopathy; however 1-hexadecanol, a defense volatile compound correlated positively with principal component (PC) 1. The chemical constitution of branches subject to exclusion plus damage or plus epiphytes was similar among them. PC2 indicated that palmitic acid (allelopathic compound) and squalene, a triterpene that attracts herbivore enemies, correlated positively with the inhibition of seed germination of T. recurvata. Inhibition of seed germination of T. recurvata was mainly correlated with the increment of palmitic acid and this compound reached higher concentrations in excluded branches treatments. Then, it is likely that the allelopathic response of I. murucoides would increase to the damage (shade, load) that may be caused by a high load of epiphytes than to damage caused by the xylophages.     

水稻化感物质抑草作用机理的分子生物学研究

水稻化感作用是通过水稻植株体向环境中释放化感物质来实现的．水稻化感物质的主要抑草作用机理有：抑制杂草种子的发芽,影响激素平衡,破坏细胞膜系统的完整性,影响光合作用和呼吸作用,干扰对营养和水分的吸收,影响蛋白质合成和基因表达等．水稻化感作用是由多基因控制的,表现为数量性状．利用分子生物学技术和化感生物检测技术等研究手段检测到了多个水稻化感作用的主效应(QTLs)位点,但不同水稻化感种质其主效应QTLs位点明显不同．通过分子辅助选择和建立近等基因系的方法对检测到的QTLs作进一步的精细定位,最终实现水稻化感抑草有利基因的克隆是今后研究的重要方向。     

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.     

高寒草场优势杂草黄帚橐吾水浸液对牧草的化感作用

研究了高寒草场中主要毒杂草——黄帚橐吾水浸液对同域分布的5种牧草的化感作用.将反应指数(RI)相加平均后分为一、二、三级敏感指数,分别评价不同测定指标、不同发育期和物种水平对化感作用的敏感性;从供体角度对相应RI值做同样处理,以综合评价不同水浸液及其不同浓度的化感效应.结果表明,5种牧草对黄帚橐吾化感作用的敏感性从强到弱依次为早熟禾>大雀麦>中华羊茅>羊茅>垂穗披碱草;牧草种子萌发期的敏感性强于幼苗生长期,尤以萌发指数受化感作用的影响明显.黄帚橐吾根、叶水浸液对牧草生物测定结果大多达到显著差异水平,物种水平的化感效应均表现为抑制作用,说明雨水淋溶是黄帚橐吾向环境中释放化感物质的途径之一.黄帚橐吾根部化感效应强于叶片,可能与其地下种间激烈的资源竞争有关.化感作用在增强黄帚橐吾生存竞争力、扩大种群和入侵草场中起着不容忽视的作用,可能是黄帚橐吾单优势种群落形成和草场退化的原因之一.     

水稻化感作用研究综述

主要对水稻化感现象发现,化感生物检测,化感物质分离和鉴定,化感种质资源等方面的研究进展了系统的综合论述,在此基础上,进一步对化感作用研究中的主要问题,例如;生物检测手段,化感作用利用途径等进行了分析。并对我国的化感作用研究的存在问题及发展方向作出了论述。     

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.