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.     

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.     

Biochemical frontiers of allelopathy

Allelopathic interactions between plants and other organisms have been recognized by scientists worldwide because they offer alternative uses in agriculture, such as decreasing our reliance on synthetic herbicides, insecticides, and nematicides for disease and insect control. The recognition of the role that allelopathy can have in producing optimum crop yields is of fundamental importance. Despite much optimism and some progress in unravelling the complexities of biochemical interactions between species, a firm foundation for the scientific rationale of the existence and function of the allelopathic phenomenon has not been developed. Allelopathic chemicals are primarily secondary products of plant metabolism which have been an enigma to plant scientists; however, they undergo a variety of reactions with plant, insect and animal species that inhibit or stimulate their growth and development. Examples of some allelochemicals and their basis of molecular and biological action are shown: interaction between the unicorn plant (Proboscidea louisianica L.) and cotton (Gossypium hirsutum L.); diterpenoid alkaloids (fromDelphinium ajacis L.) as allelochemicals; substances that occur in wheat (Tritcum aestivum) and wheat soil that cause autotoxic effects; alfalfa (Medicago sativa L.) root saponins as allelochemicals; humic acids from wheat soil as allelochemicals; and structure-function of flavonols serving as allelochemicals in chloroplast-mediated electron transport and phosphorylation. This paper concludes with a discussion of some frontier areas of research in allelopathy.     

Chemicals mediate the interaction between plant-associated fungi and insects

Plants, insects, and fungi have successfully colonized almost all terrestrial ecosystems, and their interactions have been the subject of numerous studies in recent decades. Plant-associated fungi include endophytic, arbuscular mycorrhizal, ambrosia, saprotrophic, pathogenic, and floral fungi. These fungi interact with insects through various mechanisms, including the modification of plant nutritional quality and degradation of plant defensive allelochemicals that are toxic to insects. Additionally, certain fungi assist plants in defending against insect attacks. Correspondingly, insects have evolved sophisticated nervous, digestive, and muscular systems that assist them in recognizing, preying on, and dispersing plant-associated fungi; these organ systems allow insects to detect and respond to various chemical signatures in the environment. Insects can be nourished, attracted, repelled, poisoned, and killed by chemical molecules produced by plant-associated fungi, which could be beneficial or detrimental to plants. This review summarizes the functions of different chemicals from the perspective of plant–fungus–insect interactions and discusses the challenges and future perspectives in this chemical ecology research field.     

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.     

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.     

Assessing the impact of transgenic plant products on soil organisms

Little is known about the impact of transgenic plant products on soil organisms. However, previous research with synthetic organics, allelochemicals, and extracellular enzymes can be used to guide future research in this area. Projects designed to quantify the impact of transgenic plants on soil organisms must clearly establish that the gene products are responsible for any observed changes. This can only be achieved by determining the fate of transgenic plant gene products during the period of the soil bioassay. The overall impact of transgenic plants will be dictated by not only the primary gene product, but secondary products resulting from abiotic and biotic soil reactions. Primary and secondary products may exhibit both acute and chronic impacts. Such impacts are best quantified using a soil microcosm in which fungal populations and micro- and mesofauna are monitored.     

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.     

Soil biota reduce allelopathic effects of the invasive Eupatorium adenophorum

Allelopathy has been hypothesized to play a role in exotic plant invasions, and study of this process can improve our understanding of how direct and indirect plant interactions influence plant community organization and ecosystem functioning. However, allelopathic effects can be highly conditional. For example allelopathic effects demonstrated in vivo can be difficult to demonstrate in field soils. Here we tested phytotoxicity of Eupatorium adenophorum (croftonweed), one of the most destructive exotic species in China, to a native plant species Brassica rapa both in sand and in native soil. Our results suggested that natural soils from different invaded habitats alleviated or eliminated the efficacy of potential allelochemicals relative to sand cultures. When that soil is sterilized, the allelopathic effects returned; suggesting that soil biota were responsible for the reduced phytotoxicity in natural soils. Neither of the two allelopathic compounds (9-Oxo-10,11-dehydroageraphorone and 9b-Hydroxyageraphorone) of E. adenophorum could be found in natural soils infested by the invader, and when those compounds were added to the soils as leachates, they showed substantial degradation after 24 hours in natural soils but not in sand. Our findings emphasize that soil biota can reduce the allelopathic effects of invaders on other plants, and therefore can reduce community invasibility. These results also suggest that soil biota may have stronger or weaker effects on allelopathic interactions depending on how allelochemicals are delivered.     

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.     

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.     

Dissipative structures and amplification of enantiomeric excess (an experimental point of view)

Various mathematical models have been proposed to account for the origin of chiral molecules in biological systems. Most of these models invoke non-linear phenomena, and are based on the general concept of dissipative structures. These theoretical models define the fundamental criteria which must be obeyed by the experimental systems that we have investigated. Our initial approach to this problem was an extensive search of the literature data in order to select a few systems or experimental situations which would satisfy the criteria defined by the theoretical models. For these reasons, we carried out a study of the possibility of stereospecific autocatalysis in the asymmetric polymerisation of benzofuran. Similarly, the formation of spatial dissipative structures by coupling of a transport process with an interfacial reaction was investigated as a simple experimental example of symmetry breaking.     

How genetic modification of roots affects rhizosphere processes and plant performance

Genetic modification of plants has become common practice. However, root-specific genetic modifications have only recently been advocated. Here, a review is presented regarding how root-specific modifications can have both plant internal and rhizosphere-mediated effects on aboveground plant properties and plant performance. Plant internal effects refer to pleiotropic processes such as transportation of the modified gene product. Rhizosphere-mediated effects refer to altered plant-rhizosphere interactions, which subsequently feed back to the plant. Such plant-soil feedback mechanisms have been demonstrated both in natural systems and in crops. Here how plant internal and rhizosphere-mediated effects could enhance or counteract improvements in plant properties for which the genetic modification was intended is discussed. A literature survey revealed that rice is the most commonly studied crop species in the context of root-specific transgenesis, predominantly in relation to stress tolerance. Phytoremediation, a process in which plants are used to clean up pollutants, is also often an objective when transforming roots. These two examples are used to review potential effects of root genetic modifications on shoots. There are several examples in which root-specific genetic modifications only lead to better plant performance if the genes are specifically expressed in roots. Constitutive expression can even result in modified plants that perform worse than non-modified plants. Rhizosphere effects have rarely been examined, but clearly genetic modification of roots can influence rhizosphere interactions, which in turn can affect shoot properties. Indeed, field studies with root-transformed plants frequently show negative effects on shoots that are not seen in laboratory studies. This might be due to the simplified environments that are used in laboratories which lack the full range of plant-rhizosphere interactions that are present in the field.     

Allelopathic Bacteria and Their Impact on Higher Plants

The impact of allelopathic, nonpathogenic bacteria on plant growth in natural and agricultural ecosystems is discussed. In some natural ecosystems, evidence supports the view that in the vicinity of some allelopathically active perennials (e.g., Adenostoma fasciculatum, California), in addition to allelochemicals leached from the shrub's canopy, accumulation of phytotoxic bacteria or other allelopathic microorganisms amplify retardation of annuals. In agricultural ecosystems allelopathic bacteria may evolve in areas where a single crop is grown successively, and the resulting yield decline cannot be restored by application of minerals. Transfer of soils from areas where crop suppression had been recorded into an unaffected area induced crop retardation without readily apparent symptoms of plant disease. Susceptibility of higher plants to deleterious rhizobacteria is often manifested in sandy or so-called skeletal soils. Evaluation of phytotoxic activity under controlled conditions, as well as ways to apply allelopathic bacteria in the field, is approached. The allelopathic effect may occur directly through the release of allelochemicals by a bacterium that affects susceptible plant(s) or indirectly through the suppression of an essential symbiont. The process is affected by nutritional and other environmental conditions, some may control bacterial density and the rate of production of allelochemicals. Allelopathic nonpathogenic bacteria include a wide range of genera and secrete a diverse group of plant growth-mediating allelochemicals. Although a limited number of plant growth-promoting bacterial allelochemicals have been identified, a considerable number of highly diversified growth-inhibiting allelochemicals have been isolated and characterized. Some species may produce more than one allelochemical; for example, three different phyotoxins, geldanamycin, nigericin, and hydanthocidin, were isolated from Streptomyces hygroscopicus. Efforts to introduce naturally produced allelochemicals as plant growth-regulating agents in agriculture have yielded two commercial herbicides, phosphinothricin, a product of Streptomyces viridochromogenes, and bialaphos from S. hygroscopicus. Many species of allelopathic bacteria that affect growth of higher plants are not plant specific, but some do exhibit specificity; for example, dicotyledonous plants were more susceptible to Pseudomonas putida than were monocotyledons. Differential susceptibility of higher plants to allelopathic bacteria was noted also in much lower taxonomical categories, at the subspecies level, in different cultivars of wheat, or of lettuce. Therefore, when test plants are employed to evaluate bacterial allelopathy, final evaluation must include those species that are assumed to be suppressed in nature. The release of allelochemicals from plant residues in plots of ‘continuous crop cultivation’ or from allelopathic living plants may induce the development of specific allelopathic bacteria. Both the rate by which a bacterium gains from its allelopathic activity through utilizing plant excretions, and the reasons for the developing of allelopathic bacteria in such habitats, are important goals for further research.     

Mycorrhizal networks: Mechanisms,ecology and modelling

Mycorrhizal networks, defined as a common mycorrhizal mycelium linking the roots of at least two plants, occur in all major terrestrial ecosystems. This review discusses the recent progress and challenges in our understanding of the characteristics, functions, ecology and models of mycorrhizal networks, with the goal of encouraging future research to improve our understanding of their ecology, adaptability and evolution. We focus on four themes in the recent literature: (1) the physical, physiological and molecular evidence for the existence of mycorrhizal networks, as well as the genetic characteristics and topology of networks in natural ecosystems; (2) the types, amounts and mechanisms of interplant material transfer (including carbon, nutrients, water, defence signals and allelochemicals) in autotrophic, mycoheterotrophic or partial mycoheterotrophic plants, with particular focus on carbon transfer; (3) the influence of mycorrhizal networks on plant establishment, survival and growth, and the implications for community diversity or stability in response to environmental stress; and (4) insights into emerging methods for modelling the spatial configuration and temporal dynamics of mycorrhizal networks, including the inclusion of mycorrhizal networks in conceptual models of complex adaptive systems. We suggest that mycorrhizal networks are fundamental agents of complex adaptive systems (ecosystems) because they provide avenues for feedbacks and cross-scale interactions that lead to self-organization and emergent properties in ecosystems. We have found that research in the genetics of mycorrhizal networks has accelerated rapidly in the past 5 y with increasing resolution and throughput of molecular tools, but there still remains a large gap between understanding genes and understanding the physiology, ecology and evolution of mycorrhizal networks in our changing environment. There is now enormous and exciting potential for mycorrhizal researchers to address these higher level questions and thus inform ecosystem and evolutionary research more broadly.     

Genetic variation for sensitivity to a thyme monoterpene in associated plant species

Recent studies have shown that plant allelochemicals can have profound effects on the performance of associated species, such that plants with a history of co-existence with “chemical neighbour” plants perform better in their presence compared to naïve plants. This has cast new light on the complexity of plant–plant interactions and plant communities and has led to debates on whether plant communities are more co-evolved than traditionally thought. In order to determine whether plants may indeed evolve in response to other plants’ allelochemicals it is crucial to determine the presence of genetic variation for performance under the influence of specific allelochemicals and show that natural selection indeed operates on this variation. We studied the effect of the monoterpene carvacrol—a dominant compound in the essential oil of Thymus pulegioides—on three associated plant species originating from sites where thyme is either present or absent. We found the presence of genetic variation in both naïve and experienced populations for performance under the influence of the allelochemical but the response varied among naïve and experienced plant. Plants from experienced populations performed better than naïve plants on carvacrol soil and contained significantly more seed families with an adaptive response to carvacrol than naïve populations. This suggests that the presence of T. pulegioides can act as a selective agent on associated species, by favouring genotypes which perform best in the presence of its allelochemicals. The response to the thyme allelochemical varied from negative to neutral to positive among the species. The different responses within a species suggest that plant–plant interactions can evolve; this has implications for community dynamics and stability.     

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。     

Controlled-Environment Sunlit Plant Growth Chambers

Controlled environment sunlit plant growth chambers have been built because of a great interest in plant responses to environmental variables under light intensities approaching those of natural sunlight conditions. Individual research projects have designed sunlit chambers that differ in size, structure, material, and environmental control systems dependent on the goals of the projects. Most literature describes plant organism responses to environmental variables, whereas reports of system design and performance are few. The objective of this article is to present a review of the engineering aspects of the design, environmental control, and performance of sunlit growth chambers that have been described in the literature. Most controlled environment sunlit growth chambers have been constructed with experimental plants grown in either pots or soil bins. Although a few sunlit growth chambers were designed for field grown plants, precise environmental control was not available. Further progress in the development of precise controlled environment sunlit growth chambers should include portability (or movability) so these chambers can be used in multiple field sites for greater cost-effectiveness. Modifications for improvements in guidelines for the design and operation of controlled environment growth chamber studies are also proposed.