赤潮过程中“藻-菌”关系研究进展

微生物对促进海洋物质循环,维持水生环境的生态平衡具有重要作用。在赤潮事件中,基于微生物(尤其是细菌)的多样性和重要性,它们与藻类之间的相互关系成为了研究的热点。过去20年里,人们从不同角度对"藻-菌"间的关系进行了探索,包括物理学过程、生物学过程、环境过程以及化学过程。就化学过程而言,它作为一种较早出现的技术,在以往的研究中带给人们许多认识藻菌关系的方法。随着学科的渗透,化学法有了拓展与延伸,为人们认识藻菌关系带来了新的契机。从化学生态学领域来梳理"藻-菌"关系中涉及的现象和行为,包括菌对藻的有益面、菌对藻的有害面、以及藻类应答细菌行为的化学途径;并从信号语言(群体感应、化感作用)的角度来阐释两者之间的互生或克生关系。通过文献综述的方式来解读藻菌关系的互作过程和机理,为认识赤潮的发生和防控方法提供借鉴。     

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.     

Harmful algal blooms of allelopathic microalgal species: The role of eutrophication

The ability of certain harmful algal species to produce and release chemicals that inhibit the growth of co-occurring phytoplankton species, here considered as allelopathy, is closely associated with competition for limiting nutrient resources. Many phytoplankton cells are known to release elevated amounts of organic compounds under nutrient limitation. Eutrophication alters the nitrogen-to-phosphorus balance and, when nutrient availability is unbalanced, nutrient limitation may result. Algal species that can compete successfully for available growth-limiting nutrient(s) have the potential to become dominant and form blooms. The stress conditions imposed by the shifted nutrient supply ratios can, in some algae, stimulate production of allelochemicals that inhibit potential competitors. Thus, under cultural eutrophication, altered nutrient (N, P) ratios and limiting nutrient supplies can stimulate increased production of allelochemicals, including toxins, by some algal species and accentuate the adverse effects of these substances on other algae. Future investigation on the characterization of the chemical compounds involved in the allelopathic process are needed to advance the study of the mode of action of phytoplankton allelochemicals.     

Allelopathy of Aquatic Autotrophs

Allelopathy in aquatic environments may provide a competitive advantage to angiosperms, algae, or cyanobacteria in their interaction with other primary producers. Allelopathy can influence the competition between different photoautotrophs for resources and change the succession of species, for example, in phytoplankton communities. Field evidence and laboratory studies indicate that allelopathy occurs in all aquatic habitats (marine and freshwater), and that all primary producing organisms (cyanobacteria, micro- and macroalgae as well as angiosperms) are capable of producing and releasing allelopathically active compounds. Although allelopathy also includes positive (stimulating) interactions, the majority of studies describe the inhibitory activity of allelopathically active compounds. Different mechanisms operate depending on whether allelopathy takes place in the open water (pelagic zone) or is substrate associated (benthic habitats). Allelopathical interactions are especially common in fully aquatic species, such as submersed macrophytes or benthic algae and cyanobacteria. The prevention of shading by epiphytic and planktonic primary producers and the competition for space may be the ultimate cause for allelopathical interactions. Aquatic allelochemicals often target multiple physiological processes. The inhibition of photosynthesis of competing primary producers seems to be a frequent mode of action. Multiple biotic and abiotic factors determine the strength of allelopathic interactions. Bacteria associated with the donor or target organism can metabolize excreted allelochemicals. Frequently, the impact of surplus or limiting nutrients has been shown to affect the overall production of allelochemicals and their effect on target species. Similarities and differences of allelopathic interactions in marine and freshwater habitats as well as between the different types of producing organisms are discussed. Referee: Dr. Friedrich Jüttner, Universität Zürich-Limnologische Station, Institut für Pflanzen biologie, Universität Zürich, Seestrasse 187, Ch-8802 Klichberg ZH, Switzerland     

赤潮藻毒素生物合成研究进展

合成毒素是赤潮藻类的一个常见特征,已知能够产生毒素的微藻有70多种。作为次级代谢产物,藻毒素的产生可能是一种压制或清除其它藻类竞争者的一种反应,在群落演替、种间竞争中发挥重要作用。目前,人们对藻毒素生物合成机理依然知之甚少,相关基因的研究仍无明显突破。利用环境因子诱导毒素生成变化进而分离差异表达基因或者比较不同产毒藻株间基因表达的差异,从中克隆藻毒素生物合成基因似乎是一种极具潜力的研究方向。     

马来眼子菜化感作用对斜生栅藻同工酶的影响

采用聚丙烯酰胺凝胶电泳的方法,研究了马来眼子菜(Potamogeton malaianus)和斜生栅藻(Scenedesmusobliquus)共培养系统中斜生栅藻的过氧化物酶(POD)、超歧化物氧化酶(SOD)、过氧化氢酶(CAT)、细胞色素氧化酶(COD)等同工酶的变化,探讨了沉水植物化感作用对藻类部分酶类的影响。与正常生长状态的对照组相比,马来眼子菜作用下的样品组斜生栅藻POD酶带总数保持稳定,但酶谱组成发生变化;样品组斜生栅藻SOD酶带数量减少而CAT酶带数量增加。综合结果表明,样品组藻体在化感作用下通过改变酶活性、调整同工酶组成等来抵御化感作用带来的活性氧(ROS)伤害。马来眼子菜作用下的样品组藻体COD同工酶的酶谱变化明显,说明化感作用在一定程度上影响了斜生栅藻的生物氧化过程。     

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。     

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.     

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.     

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。     

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.     

Toxin-assisted micropredation: experimental evidence shows that contact micropredation rather than exotoxicity is the role of Prymnesium toxins

Blooms of Prymnesium parvum can severely harm fish and zooplankton, presumably through the release of allelopathic exotoxins that offer advantages for Prymnesium in its interactions with competitors and prey. We show that Prymnesium attaches to zooplankton and fish, causing mortality, whereas exposure of these organisms to Prymnesium across a permeable membrane does not cause mortality. We also show that Prymnesium exotoxins are released independently of contact toxicity only in response to experimental procedures or natural causes of stress. Our results are consistent with the idea that toxins have evolved for release during cell-to-cell contact in support of heterotrophy. The evolution of toxin-assisted micropredation would be consistent with mechanisms of natural selection favouring individual fitness as opposed to broadcast allelopathy from which the benefits are more dispersed. Research into the toxicity of Prymnesium and other harmful algal species may profit from focus on processes following physical contact with potential prey.     

数学建模在生物化感及其赫米西斯(Hormesis)现象中的应用

简述了国内外对生物化感数学建模的研究近况,包括通过数学模型描述化感物质的赫米西斯(Hormesis)现象,化感作用在受体植物不同密度条件下的表达,植物残茬中化感物质的分解动态及受体植物的动态响应,环境中化感物质的动态变化规律及在植物-昆虫-天敌系统中的应用等。并对化感数学建模领域的先驱机理模型An-Hormesis模型,An-Liu-Johnson-Lovett模型,和An_Residue模型做了简介。     

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。     

Competition and allelopathy in aquatic plant communities

The paper reviews the published literature on the studies of competition and allelopathy in aquatic plant communities. Taking a broader view of the community, the studies on interactions between macrophytes and microphytes, macrophytes and macro-invertebrates and microbial communities are also reviewed. The role of these interactions in the structure and dynamics of aquatic communities has been discussed in light of the current hypotheses concerning competition in terrestrial communities. The available information suggests that the aquatic plants of various growth forms differ greatly among themselves in their responses and adaptations to competition and allelopathy. The possible application of these interactions in biological control of plant pests and in agriculture is also summarized. We conclude that the observed differences in these interactions between the terrestrial and aquatic environment are due to the effects of water as a non-resource variable as well as due to special adaptive characteristics of aquatic plants. Further we hypothesize that the aquatic plants adopt both competitive and allelopathic strategies under different conditions and in interactions with different plants. The review highlights that our knowledge of both competition and allelopathy among aquatic plant communities is inadequate and fragmentary, and therefore, both extensive and intensive studies are required.     

入侵植物北美车前和野胡萝卜种内及种间的化感作用

当前入侵植物种类和入侵范围均日益扩大,而多种入侵植物共同入侵同一生境的概率也逐渐增加。为探讨这些入侵植物种内和种间的化感作用以及化感作用对其竞争能力的影响,本研究以北美车前(Plantago virgica)和野胡萝卜(Daucus carota)为例,采用室内生物学测定法,分析了2种植物不同器官(根、茎或花序轴、叶)、不同浓度(CK、0.005、0.01和0.05 g·mL^-1)的水浸提液培养对其自身和对方种子萌发和幼苗生长的影响。结果表明,浸提液浓度对2种入侵植物种子萌发和幼苗生长的影响最大,且浓度、器官及物种之间存在显著交互作用。2物种各器官各浓度浸提液对二者种子萌发总体表现为抑制;中低浓度浸提液对同种植物幼苗生长有一定促进作用;高浓度浸提液尤其是野胡萝卜叶浸提液对二者种子萌发与幼苗生长均有显著抑制作用。综合化感效应指数表明,浸提液浓度越高抑制作用越强,地上部分化感效应强于地下,且野胡萝卜对北美车前的种间化感作用最强。由此可见,种内和种间化感作用是入侵植物在相同入侵地相互竞争的重要方式。     

Does allelopathy offer real promise for practical weed management and for explaining rhizosphere interactions involving higher plants?

This paper selectively reviews developments in the study of allelopathy, principally from a chemical standpoint. Particular attention is given to recently discovered aspects of signalling within the rhizosphere rather than to direct physiological effects caused by broadly active allelopathic agents with only a brief review of the role of bulk generation of weakly biocidal material. Many of the chemicals responsible for allelopathic effects have been identified and the biosynthesis and modes of action of a number are included. The paper also highlights instances where signalling aspects of allelopathy may be exploited for practical weed control purposes, and how the study of allelopathic components of the rhizosphere metabolome might be studied more directly by means of plant genomics.     

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.