Searching for allelopathic effects of submerged macrophytes on phytoplankton—state of the art and open questions

Allelopathy, here defined as biochemical interactions between aquatic primary producers, has always been intriguing as a process explaining the dominance of certain plant or algal species over others. Negative chemical interference has been invoked as one of the steering mechanisms behind mutual dominance of either submerged macrophytes or phytoplankton in shallow eutrophic lakes. Yet, despite much effort, convincing evidence for allelopathic interactions in situ is still missing. Also, laboratory approaches often lack reality. Inspired by a series of talks at the Shallow Lakes 2005 meeting in Dalfsen, the Netherlands, we argue that there is circumstantial but strong evidence that allelopathic interference between submerged macrophytes and phytoplankton may indeed exist in aquatic ecosystems despite the problems associated with research in this field. We first discuss experimental approaches combining laboratory and field studies, based on examples presented at this meeting. We then discuss the impact of nutrient status of both producing and target organism and biotic factors such as herbivory or pathogens that might affect allelopathy. Further topics are the potential seasonality of effects and the species-specificity of certain allelochemicals. We conclude with some thoughts why a final proof for allelopathy in situ might remain difficult or even inaccessible in some cases, and why we nevertheless should not abandon this idea.     

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。     

Can allelopathically active submerged macrophytes stabilise clear-water states in shallow lakes?

Inhibition of phytoplankton by allelochemicals released by submerged macrophytes is supposed to be one of the mechanisms that contribute to the stabilisation of clear-water states in shallow lakes. The relevance of this process at ecosystem level, however, is debated because in situ evidence is difficult to achieve. Our literature review indicates that allelopathically active species such as Myriophyllum, Ceratophyllum, Elodea and Najas or certain charophytes are among the most frequent submerged macrophytes in temperate shallow lakes. The most common experimental approach for allelopathic interference between macrophytes and phytoplankton has been the use of plant extracts or purified plant compounds. Final evidence, however, requires combination with more realistic in situ experiments. Such investigations have successfully been performed with selected species. In situ allelopathic activity is also influenced by the fact that phytoplankton species exhibit differential sensitivity against allelochemicals both between and within major taxonomic groups such as diatoms, cyanobacteria and chlorophytes. In general, epiphytic species apparently are less sensitive towards allelochemicals than phytoplankton despite living closely attached to the plants and being of key importance for macrophyte growth due to their shading. Light and nutrient availability potentially influence the sensitivity of target algae and cyanobacteria. Whether or not additional stressors such as nutrient limitation enhance or dampen allelopathic interactions still has to be clarified. We strongly propose allelopathy as an important mechanism in the interaction between submerged macrophytes and phytoplankton in shallow lakes based on the frequent occurrence of active species and the knowledge of potential target species. The role of allelopathy interfering with epiphyton development is less well understood. Including further levels of complexity, such as nutrient interference, grazing and climate, will extend this ecosystem-based view of in situ allelopathy.     

An ecological perspective of allelochemical interference in land–water interface communities

Allelochemical interactions among aquatic macrophytes and between macrophytes and attached microbial assemblages (epiphyton) influence a number of ecological processes. The ecological importance of these interactions, however, is poorly understood; we hypothesize that paucity has resulted, in part, from (1) a narrow focus on exploration for herbicidal plant products from aquatic macrophytes, (2) the difficulties in distinguishing resource competition from allelopathic interference, and (3) a predominance of approaching aquatic allelopathy from a terrestrial perspective. Based upon recent thorough investigations of allelopathy among aquatic vascular plants, chemical compounds that influence competitive interactions among littoral organisms are amphiphilic compounds that tend to remain near the producing organism (e.g., polyphenolic compounds and volatile fatty acids). Production of these compounds may be influenced by relative availability of nutrients (particularly phosphorus and nitrogen), inorganic carbon, and light. Macrophyte strategies of clonal reproduction, in an effort to persist in these highly productive and competitive habitats, have contributed to reduced reliance upon sexual reproduction that is correlated with allelopathic autotoxicity among several dominant wetland plant species. Although few studies document the importance of allelochemical interactions in the wetland and littoral zones of aquatic ecosystems, abundant evidence supports the potential for significant effects on competition and community structure; effects of altered nutrient ratios and availability on plant chemical composition; and resultant effects on trophic interactions, particularly suppression of herbivory, competitive attached algae and cyanobacteria, and heterotrophic utilization of organic matter by bacteria and fungi.     

Algal allelopathy

The comprehensive review on allelopathy (Rice, 1979, 1984) has been largely responsible for the evolution of allelopathy as an independent branch of chemical/ physiological ecology. The allelopathic research during the last four decades drew attention to different facets of the interactions among the constituents of habitat, calling for an understanding of the role of allelopathy under different habitat conditions. In view of this, we have reviewed the existing information on allelopathic interactions in aquatic habitats with special reference to algal allelopathy. This review has been mainly confined, therefore, to different aspects of algal allelopathy such as allelopathic interactions in algae, algal toxins, bioassays, and implications of algal allelopathy. In spite of the large number of reviews on allelopathy (see section III), no independent review appears on algal allelopathy. Although there were reports of toxins from cyanobacteria and other algae, no appreciable attempt was made to implicate algal toxins in allelopathy under field conditions. Knowledge of chemistry and biology of allelochemical can help in their potential use in controlling plant diseases and weeds. Therefore, it is urgent to study algal toxins for their involvement in ecological phenomena such as succession, for their uses as herbicides, weedicides, and pesticides, for their uses in solving some of the problems of algal ecology, and for their involvement in applied aspects.     

Allelopathic agents from aquatic ecosystems: potential biopesticides models

Allelopathy is usually regarded as a terrestrial science and little attention has been paid to the allelochemicals involved in the interactions between aquatic autotrophs. Also, chemicals involved in the interaction between algae (macro or microscopic) and their environment are not often acknowledged as allelopathic. Allelopathy is a key factor for explaining microalgae assemblages, community structure and the dynamics of the populations within the ecosystem. Herein we present an overview of what is known in this field, describing the chemicals yet identified and which kind of interactions are they involved in.     

淡水水生植物化感作用研究进展

水生植物的化感作用对淡水生态系统中水草的可持续管理和湖泊富营养化的生态控制具有非常重要的意义。本文综述了淡水水生植物化感作用的发展历史和研究现状,讨论了水生植物化感作用研究过程中所涉及的实验方法、生物测试方法、化感物质分离方法以及影响水生植物化感研究的环境因素。抑藻圈试验、藻类生长试验和浮萍生长试验是常用的生物测试方法;采用共生培养或单独培养的方法,从种植水中分离、鉴定化感物质,对证实复杂水环境中水生植物化感作用的存在,研究它们化感物质的释放途径和作用机理都具有十分重要的作用。     

植物间的相生相克效应

本文介绍了相生相克的定义,相互抑制和刺激促进现象,作用的影响因素,相生相克化合物种类等。首次对其化合物的提取、分离鉴定和生物测试等方法进行了较全面的叙述。从总体上对相生相克作用进行了较系统的评述。     

植物间的相生相克效应

本文介绍了相生相克的定义,相互抑制和刺激促进现象,作用的影响因素,相生相克化合物种类等。首次对其化合物的提取、分离鉴定和生物测试等方法进行了较全面的叙述。从总体上对相生相克作用进行了较系统的评述。     

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.     

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.     

Allelopathic activity of Stratiotes aloides on phytoplankton—towards identification of allelopathic substances

The allelopathic activity of the aquatic macrophyte, Stratiotes aloides, was determined with laboratory experiments. Active compounds exuded in the medium or present in plant tissue were extracted using standard procedures and solid phase extraction (SPE). The activity towards various cyanobacteria and chlorophytes was tested in two different bioassay systems using agar plates and liquid cultures of phytoplankton. Extracts and exudates of S. aloides affected phytoplankton growth. SPE-enriched exudates and enriched water from a natural Stratiotes stand caused inhibition of target species, however, also some controls were active. Phytoplankton species exhibited differential sensitivity to extracts of S. aloides. We observed inhibitory and stimulatory effects on phytoplankton. In general, more cyanobacteria than other phytoplankton species were inhibited, and the inhibition of cyanobacteria was stronger. In most cases, nutrient (P or K) limitation of Synechococcus elongatus and Scenedesmus obliquus decreased the sensitivity of these species towards allelochemicals from Stratiotes aloides, except for P-limited cultures of Scenedesmus. The allelopathically active compound(s) from Stratiotes are moderately lipophilic and most likely no phenolic compounds. Our results indicate that allelopathy (besides nutrient interference and shading) might also account for the low phytoplankton and filamentous algae densities in the vicinity of Stratiotes plants, at least during certain phases of the life-cycle of Stratiotes.     

化感作用的研究意义及发展前景

对化感作用在生态学、林业、农业等领域的理论及实践意义进行综述与探讨,并分析了其今后的发展方向.化感作用有助于更加合理地解释生态系统中植物组成与分布、群落演替、协同进化和生物入侵等现象;在林业生产的森林更新、混交林培育中有时会起到决定性作用;在农业上,单作、轮作、覆盖等各种种植方式均受化感影响,此外,一些化感物质可用作杀虫剂和除草剂,从而减轻对环境的污染水体中同样存在化感作用.培育新的抗虫害、抑草品种,分离、鉴定新的化感物质,深入揭示化感作用的机制,是今后化感作用的研究重点.     

沉水植物化感作用控藻能力评述

沉水植物所释放的化学物质对藻类的抑制作用是浅水湖泊维持清水状态的机制之一.本文从具有化感活性的沉水植物在湖泊中出现的频度、盖度、化感物质的种类、抑藻效应等方面对化感控藻进行了评述.已有研究结果表明：穗花狐尾藻、金鱼藻、伊乐藻等是具有很高活性的沉水植物,尤其是在其生物量达到一定程度,且湖泊中的优势藻为较敏感的种类时,沉水植物分泌的化感物质对浮游藻类的抑制作用更强;沉水植物释放的多酚类等化感物质具有控藻能力;化感物质对于不同种类藻的抑制作用具有选择性,蓝藻和硅藻比绿藻更为敏感,附生藻类通常比浮游藻类具有更高的耐受性;环境因素如光照、营养限制、温度等会显著影响沉水植物化感作用效果.沉水植物的化感控藻研究尚处于初始阶段,关于环境因素对化感作用的影响、化感物质的分离鉴定、选择性抑藻机理以及化感物质代谢途径等方面还有待深入、全面的研究.     

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.     

Allelopathic interactions between plants. Multi site action of allelochemicals

Allelopathy is defined as mechanism of plant-plant, plant-microorganisms, plant-virus, plant-insect, and plant-soil-plant interactions mediated by plant- or microorganism-produced chemicals released to the environment. The majority of allelochemicals are secondary metabolites and among others belong to terpenoids, phenolic compounds, organic cyanides and longchain fatty acids. The action of allelochemicals in target plant is diverse and affects a large number of biochemical reactions resulting in modifications of different physiological functions. Thus the results of allelochemical action can be detected at different levels of plant organization: molecular, structural, biochemical, physiological and ecological. Enzyme activities, cell division and ultrastructure, membrane permeability, ion uptake and as a consequence plant growth and development are modified by allelochemicals. Significant effects on photosynthesis and respiration are the best-characterized results of allelopathic interactions. Moreover allelopathic compounds seem to induce a secondary oxidative stress expressed as enhanced free radical production and induction of cellular antioxidant system. Plant survival under allelopathy stress conditions depends on plant defense leading to allelochemical detoxication, the process which may go on in parallel to cell defense reaction to oxidative stress. The article presents some aspects of the current knowledge regarding mechanisms of the allelopathy phenomenon. The allelopathy is a complex problem, thus comprehensive understanding of allelochemical mode of action requires further investigation and still remains an open question.     

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.     

环境胁迫下植物的化感作用及其诱导机制

植物化感作用是生态学研究中一个十分活跃的领域,对植物为什么和在什么条件下释放化感物质这一重要问题有不同的认识。在对环境胁迫下植物化感作用的变化及环境胁迫因子对化感物质听诱导机制等方面进行了评述后,指出植物化感化质的产生和释放是植物在环境胁迫的选择压力下形成的,植物化感作用是植物在进化过程中产生的一种对环境的适应性机制。     

分子水平和土壤系统化感作用研究现状与展望

化感作用是生态学研究中的十分活跃的领域之一。除概述化感作用对于植物细胞、组织以及环境胁迫对于化感效应的影响外 ,本文主要讨论了在分子和土壤生态系统水平上的化感作用的研究。在分子水平化感作用研究中 ,主要从化感基因的定位、蛋白质和核酸合成、基因表达和调控以及生化机制等几个方面进行了论述。在化感作用对于土壤生态系统的影响方面 ,主要对其在土壤性质、土壤微生物和土壤动物方面的效应进行了评述。最后 ,并就化感作用研究中存在的问题和未来发展提出一些看法与展望。