植物次生代谢及其与环境的关系

人类对植物次生代谢产物（天然产物）的早期研究源于它们的应用价值,近些年来人们越来越认识到植物次生代谢产物广泛的生物学效应,开始重新评价这些化合物在植物生命活动以及生态系统中可能扮演的角色。植物的次生代谢是植物在长期进化中与环境（生物的和非生物的）相互作用的结果,次生代谢产物在植物提高自身保护和生存竞争能力、协调与环境关系上充当着重要的角色。介绍了植物次生代谢及其产物的特点,概述了植物次生代谢与温度、水分、光照、养分、CO2浓度、UV-B辐射、环境污染等非生物环境以及与化学防御、化感作用、菌根共生、微生物病害的关系。研究植物次生代谢与环境的关系,可以从更深的层次发掘植物与环境的内在联系,为全面、深入认识植物与环境的相互关系提供新的研究途径,同时也有利于人类更有效、合理地利用植物的次生代谢产物。     

Relative importance of competition and plant–soil feedback,their synergy,context dependency and implications for coexistence

Plants interact simultaneously with each other and with soil biota, yet the relative importance of competition vs. plant–soil feedback (PSF) on plant performance is poorly understood. Using a meta‐analysis of 38 published studies and 150 plant species, we show that effects of interspecific competition (either growing plants with a competitor or singly, or comparing inter‐ vs. intraspecific competition) and PSF (comparing home vs. away soil, live vs. sterile soil, or control vs. fungicide‐treated soil) depended on treatments but were predominantly negative, broadly comparable in magnitude, and additive or synergistic. Stronger competitors experienced more negative PSF than weaker competitors when controlling for density (inter‐ to intraspecific competition), suggesting that PSF could prevent competitive dominance and promote coexistence. When competition was measured against plants growing singly, the strength of competition overwhelmed PSF, indicating that the relative importance of PSF may depend not only on neighbour identity but also density. We evaluate how competition and PSFs might interact across resource gradients; PSF will likely strengthen competitive interactions in high resource environments and enhance facilitative interactions in low‐resource environments. Finally, we provide a framework for filling key knowledge gaps and advancing our understanding of how these biotic interactions influence community structure.     

Differences in Competitive Ability between Plants from Nonnative and Native Populations of a Tropical Invader Relates to Adaptive Responses in Abiotic and Biotic Environments

The evolution of competitive ability of invasive plant species is generally studied in the context of adaptive responses to novel biotic environments (enemy release) in introduced ranges. However, invasive plants may also respond to novel abiotic environments. Here we studied differences in competitive ability between Chromolaena odorata plants of populations from nonnative versus native ranges, considering biogeographical differences in both biotic and abiotic environments. An intraspecific competition experiment was conducted at two nutrient levels in a common garden. In both low and high nutrient treatments, C. odorata plants from nonnative ranges showed consistently lower root to shoot ratios than did plants from native ranges grown in both monoculture and competition. In the low nutrient treatment, C. odorata plants from nonnative ranges showed significantly lower competitive ability (competition-driven decreases in plant height and biomass were more), which was associated with their lower root to shoot ratios and higher total leaf phenolic content (defense trait). In the high nutrient treatment, C. odorata plants from nonnative ranges showed lower leaf toughness and cellulosic contents (defense traits) but similar competitive ability compared with plants from native ranges, which was also associated with their lower root to shoot ratios. Our results indicate that genetically based shifts in biomass allocation (responses to abiotic environments) also influence competitive abilities of invasive plants, and provide a first potential mechanism for the interaction between range and environment (environment-dependent difference between ranges).     

Optimal defence, kin conflict and the distribution of furanocoumarins among offspring of wild parsnip

The factors influencing the allocation of chemical defences to plant offspring have largely been unexplored, conceptually and experimentally. Because evolutionary interactions between maternal plants and their progeny can affect resource allocation patterns among sibling offspring, we suggest that kin conflict as well as herbivore–plant interaction theories need to be considered to predict chemical defence allocation patterns. Optimal defence theory predicts that maternal plants should defend more heavily those offspring in which resources have been disproportionately invested. In contrast, kin conflict theory predicts that natural selection will favour genotypes that can compete successfully for maternal defences irrespective of their quality, even at the expense of the fitness of siblings and the maternal plant. Evidence for these defence patterns were evaluated by examining the allocation of furanocoumarins to seeds of the wild parsnip (Pastinaca sativa, Apiaceae). Furanocoumarins are toxins that are localized within the oil tubes of the maternal tissues of seeds. We evaluated the role of offspring investment (endosperm mass) and seed genotype on furanocoumarin allocation by mating an array of pollen donors with pollen recipients. Furanocoumarins were found to be positively correlated with endosperm mass on one side of the seed, a result consistent with optimal defence theory; however, on the other side of the seed, furanocoumarin content was influenced by seed genotype and was unrelated to endosperm mass. These effects varied with maternal plant. Further experiments demonstrated that nearly 80% of furanocoumarin production occurs after pollination, when fertilization products are active. Although the amount of furanocoumarin influenced by the seed genotype is small relative to the total quantity in the seed, these furanocoumarins are the first line of defence against important predators, such as the parsnip webworm, Depressaria pastinacella (Lepidoptera: Oecophoridae). We found that parsnip webworm larvae were able to discriminate among genotypes within an inflorescence. In line with previous studies, these results suggest that a genotype's ability to influence furanocoumarin defence may affect its probability of survival. We conclude that the distribution of defences among plant offspring in wild parsnip is probably influenced by competition among seed genotypes that conflicts with maternal optimal defence. This revised version was published online in July 2006 with corrections to the Cover Date.     

Future directions in the ontogeny of plant defence: understanding the evolutionary causes and consequences

Plant defence often varies by orders of magnitude as plants develop from the seedling to juvenile to mature and senescent stages. Ontogenetic trajectories can involve switches among defence traits, leading to complex shifting phenotypes across plant lifetimes. While considerable research has characterised ontogenetic trajectories for now hundreds of plant species, we still lack a clear understanding of the molecular, ecological and evolutionary factors driving these patterns. In this study, we identify several non‐mutually exclusive factors that may have led to the evolution of ontogenetic trajectories in plant defence, including developmental constraints, resource allocation costs, multi‐functionality of defence traits, and herbivore selection pressure. Evidence from recent physiological studies is highlighted to shed light on the underlying molecular mechanisms involved in the regulation and activation of these developmental changes. Overall, our goal is to promote new research avenues that would provide evidence for the factors that have promoted the evolution of this complex lifetime phenotype. Future research focusing on the questions and approaches identified here will advance the field and shed light on why defence traits shift so dramatically across plant ontogeny, a widespread but poorly understood ecological pattern.     

An ecophysiological approach to modelling resource fluxes in competing plants

A conceptual model of resource acquisition and allocation within a generalized, individual plant growing vegetatively in competition with others is presented. The model considers C and N acquisition, synthesis of assimilates and their transport and partitioning, growth of new tissues, reserve formation and recycling, and losses due to root exudation and respiration. These processes are regulated by the relative size of the C and N substrate pools in shoot and roots, in relation to meristematic sink strength. Translocation and allocation patterns are represented according to the Minchin phloem transport model. The current model is used to consider the impact of competition on resource acquisition and allocation, first by considering a plant growing in isolation and its response to manipulation of light, CO2 and N supplies. Secondly, competitive plants are introduced and the direct effects on plant responses in terms of resource depletion are considered separately from indirect effects such as potential changes in the quality of resources available (e.g. light quality or soil N sources). In the past, many studies of plant competition have not established the importance of these indirect effects because they have not established all the processes involved in competition. This model can be used to interpret responses of whole plants to their neighbours in terms of the relative importance of both the direct and indirect effects of competition.     

The evolution of growth rate, resource allocation and competitive ability in seeder and resprouter tree seedlings

Many woody plant species in fire disturbed communities survive disturbance events by resprouting. The resprouting life history is predicted to be costly to plants as resources are diverted into storage for post-fire regrowth rather than allocated to current growth, and resprouting species typically grow more slowly than seeder species (species that do not resprout after disturbance events). Differences in allocation to current growth are also predicted to make resprouter species poorer competitors compared to seeder species. We tested the predictions that the evolution of a resprouter life history is associated with slow growth, increased allocation to storage, and low competitive ability in woody plant seedlings. We grew eight phylogenetically independent pairs of seeder and resprouter species in competition and no competition treatments in a field experiment near Sydney, Australia. The presence of competitors reduced plant growth rates across taxa and fire response life histories. However, relative to seeder species, resprouter species were not slower growing, they did not allocate more resources to storage, and they did not have lower competitive abilities. We propose that differences in resource allocation to storage are not responsible for differences in growth rate and competitive ability. Rather, growth rate and competitive ability in seedlings are associated with key aspects of plant life history such as life-span and body size at maturity. These traits that are sometimes, but not always, related to fire response life histories.     

Plant life history and above–belowground interactions: missing links

The importance of above–belowground interactions for plant growth and community dynamics became clear in the last decades, whereas the numerous studies on plant life history improved our knowledge on eco‐evolutionary dynamics. However, surprisingly few studies have linked both research fields despite their potential to increase our mechanistic understanding of how above belowground interactions are governed. Here I briefly review studies on above–belowground interactions and plant life history and identify important research gaps. To advance our understanding of ecological strategies and eco‐evolutionary dynamics of plants and their associated organisms it is warranted to elucidate the interconnectivity and tradeoffs of plant life history traits of growth, defence, reproduction, nutrient cycling and the functional composition of above‐ and belowground heterotrophic communities. Using the concept of tradeoffs in growth, reproduction and defence we can postulate that plants in rich soil grow, reproduce and die fast whilst avoiding above‐ and belowground antagonists, whereas plants in poor soil grow slow, live and reproduce longer and invest in above‐ and belowground mutualists and defences. However, alternative scenarios are possible and depend on the selection pressure by above‐ and belowground mutualists and antagonists during plant ontogeny and via after‐life effects. To elucidate missing links between life history traits and above–belowground interactions, complementary modelling and empirical studies are needed that reveal the coupling between below‐ and aboveground plant traits of growth, defence and reproduction, their heritability and their cost/benefit relation. These cost/benefit analyses of defence should span from individuals to future generations, taking feedback effects via altered biotic communities and resource competition into account. The role of soil fertility in steering plant life history traits requires explicit testing of trans‐generational trait shifts in growth, defence, reproduction, cost/benefit of associations with mutualists and antagonists and soil feedbacks across plant genotypes/species with distinct life history traits, grown across soil fertility gradients.     

Metal complexes increase uptake of Zn and Cu by plants: implications for uptake and deficiency studies in chelator-buffered solutions

The allocation of resources among roots and shoots represents the largest flux of resources within a plant and therefore should have been selected to maximize benefits to plants. Yet, it is unclear why some species like temperate grasses have such high root length density (RLD). Either the slow rate of diffusion of inorganic N in soils or interplant competition could explain the high RLD of temperate grasses. Using a fine-scale model of nutrient dynamics in the soil and plant growth, a cost–benefit approach was used to assess optimal allocation rates for plants that accounted for value of both carbon and nitrogen. In the absence of interplant competition, resource benefits are maximized with very little root length except in extremely dry soils for ammonium. In the presence of a competitor, optimal allocation of N to roots is much greater and increases as ability of competitors to produce root length increase. Competition for inorganic nitrogen generates a classic aspect of the tragedy of the commons, the “race for fish”, where plants must allocate more resources to acquisition of the limiting resource than is optimal for plants in the absence of competition. As such, nutrient competition needs to be directly addressed when understanding plant- and ecosystem-level resource fluxes as well as the evolution of root systems.     

Interactions between functionally diverse fungal mutualists inconsistently affect plant performance and competition

Plants form mutualistic relationship with a variety of belowground fungal species. Such a mutualistic relationship can enhance plant growth and resistance to pathogens. Yet, we know little about how interactions between functionally diverse groups of fungal mutualists affect plant performance and competition. We experimentally determined the effects of interaction between two functional groups of belowground fungi that form mutualistic relationship with plants, arbuscular mycorrhizal (AM) fungi and Trichoderma, on interspecific competition between pairs of closely related plant species from four different genera. We hypothesized that the combination of two functionally diverse belowground fungal species would allow plants and fungi to partition their symbiotic relationships and relax plant–plant competition. Our results show that: 1) the AM fungal species consistently outcompeted the Trichoderma species independent of plant combinations; 2) the fungal species generally had limited effects on competitive interactions between plants; 3) however, the combination of fungal species relaxed interspecific competition in one of the four instances of plant–plant competition, despite the general competitive superiority of AM fungi over Trichoderma. We highlight that the competitive outcome between functionally diverse fungal species may show high consistency across a broad range of host plants and their combinations. However, despite this consistent competitive hierarchy, the consequences of their interaction for plant performance and competition can strongly vary among plant communities.     

The plant nitrogen mobilization promoted by Colletotrichum lindemuthianum in Phaseolus leaves depends on fungus pathogenicity

Nitrogen plays an essential role in the nutrient relationship between plants and pathogens. Some studies report that the nitrogen-mobilizing plant metabolism that occurs during abiotic and biotic stress could be a 'slash-and-burn' defence strategy. In order to study nitrogen recycling and mobilization in host plants during pathogen attack and invasion, the Colletotrichum lindemuthianum/Phaseolus vulgaris interaction was used as a model. C. lindemuthianum is a hemibiotroph that causes anthracnose disease on P. vulgaris. Non-pathogenic mutants and the pathogenic wild-type strain were used to compare their effects on plant metabolism. The deleterious effects of infection were monitored by measuring changes in chlorophyll, protein, and amino acid concentrations. It was shown that amino acid composition changed depending on the plant-fungus interaction and that glutamine accumulated mainly in the leaves infected by the pathogenic strain. Glutamine accumulation correlated with the accumulation of cytosolic glutamine synthetase (GS1 alpha) mRNA. The most striking result was that the GS1 alpha gene was induced in all the fungus-infected leaves, independent of the strain used for inoculation, and that GS1 alpha expression paralleled the PAL3 and CHS defence gene expression. It is concluded that a role of GS1 alpha in plant defence has to be considered.     

Reproductive efficiency and shade avoidance plasticity under simulated competition

Plant strategy and life‐history theories make different predictions about reproductive efficiency under competition. While strategy theory suggests under intense competition iteroparous perennial plants delay reproduction and semelparous annuals reproduce quickly, life‐history theory predicts both annual and perennial plants increase resource allocation to reproduction under intense competition. We tested (1) how simulated competition influences reproductive efficiency and competitive ability (CA) of different plant life histories and growth forms; (2) whether life history or growth form is associated with CA; (3) whether shade avoidance plasticity is connected to reproductive efficiency under simulated competition. We examined plastic responses of 11 herbaceous species representing different life histories and growth forms to simulated competition (spectral shade). We found that both annual and perennial plants invested more to reproduction under simulated competition in accordance with life‐history theory predictions. There was no significant difference between competitive abilities of different life histories, but across growth forms, erect species expressed greater CA (in terms of leaf number) than other growth forms. We also found that shade avoidance plasticity can increase the reproductive efficiency by capitalizing on the early life resource acquisition and conversion of these resources into reproduction. Therefore, we suggest that a reassessment of the interpretation of shade avoidance plasticity is necessary by revealing its role in reproduction, not only in competition of plants.     

Clonal integration beyond resource sharing: implications for defence signalling and disease transmission in clonal plant networks

Resource sharing between ramets of clonal plants is a well-known phenomenon, which allows stoloniferous and rhizomatous species to internally translocate water, mineral nutrients and carbohydrates from sites of high supply to sites of high demand. The mechanisms and implications of resource integration in clonal plants have extensively been studied in the past. Vascular ramet connections are likely to provide an excellent means to share substances other than resources, such as systemic defence signals and pathogens. The aim of this paper is to propose the idea that physical ramet connections of clonal plants can be used (1) to transmit signals, which enable members of clonal plant networks to share information about their biotic and abiotic environments, and (2) to facilitate the internal distribution of systemic pathogens in clonal plant networks and populations. We will focus on possible mechanisms as well as on potential ecological and evolutionary implications of clonal integration beyond resource sharing. More specifically, we will explore the role of physiological integration in clonal plant networks for the systemic transmission of direct and indirect defence signals after localized herbivore attack. We propose that sharing defence induction signals among ramets may be the basis for an efficient early warning system, and it may allow for effective indirect defence signalling to herbivore enemies through a systemic release of volatiles from entire clonal fragments. In addition, we will examine the role of clonal integration for the internal spread of systemic pathogens and pathogen defence signals within clonal plants. Clonal plants may use developmental mechanisms such as increased flowering and clone fragmentation, but also specific biochemical defence strategies to fight pathogens. We propose that clonal plant networks can act as stores and vectors of diseases in plant populations and communities and that clonal life histories favour the evolution of pathogens with a low virulence.     

A dynamical model of environmental effects on allocation to carbon-based secondary compounds in juvenile trees

BACKGROUND AND AIMS: Patterns and variations in concentration of carbon-based secondary compounds in plant tissues have been explained by means of different complementary and, in some cases, contradictory plant defence hypotheses for more than 20 years. These hypotheses are conceptual models which consider environmental impacts on plant internal demands. In the present study, a mathematical model is presented, which converts and integrates the concepts of the 'Growth-Differentiation Balance' hypothesis and the 'Protein Competition' model into a dynamic plant growth model, that was tested with concentration data of polyphenols in leaves of juvenile apple, beech and spruce trees. The modelling approach is part of the plant growth model PLATHO that considers simultaneously different environmental impacts on the most important physiological processes of plants. METHODS: The modelling approach for plant internal resource allocation is based on a priority scheme assuming that growth processes have priority over allocation to secondary compounds and that growth-related metabolism is more strongly affected by nitrogen deficiency than defence-related secondary metabolism. KEY RESULTS: It is shown that the model can reproduce the effect of nitrogen fertilization on allocation patterns in apple trees and the effects of elevated CO(2) and competition in juvenile beech and spruce trees. The analysis of model behaviour reveals that large fluctuations in plant internal availability of carbon and nitrogen are possible within a single vegetation period. Furthermore, the model displays a non-linear allocation behaviour to carbon-based secondary compounds. CONCLUSIONS: The simulation results corroborate the underlying assumptions of the presented modelling approach for resource partitioning between growth-related primary metabolism and defence-related secondary metabolism. Thus, the dynamical modelling approach, which considers variable source and sink strengths of plant internal resources within different phenological growth stages, presents a successful translation of existing concepts into a dynamic mathematical model.     

Fitness costs associated with acetyl‐coenzyme A carboxylase mutations endowing herbicide resistance in American sloughgrass (Beckmannia syzigachne Steud.)

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Growth and Parasite Defence in Plants; the Balance between Resource Sequestration and Retention: In Lieu of a Guest Editorial

Abstract: A hypothesis on regulation of the balance between growth and parasite defence in plants is formulated, namely that plants regulate their resource allocation in a way where stress tolerance and resistance inherently lead to constraints on growth and competitiveness. Seven reviews and the subsequent article in this issue of Plant Biology contributing to this problem are briefly introduced in context.     

Plant structural traits and their role in anti-herbivore defence

We consider the role that key structural traits, such as spinescence, pubescence, sclerophylly and raphides, play in protecting plants from herbivore attack. Despite the likelihood that many of these morphological characteristics may have evolved as responses to other environmental stimuli, we show that each provides an important defence against herbivore attack in both terrestrial and aquatic ecosystems. We conclude that leaf-mass–area is a robust index of sclerophylly as a surrogate for more rigorous mechanical properties used in herbivory studies. We also examine herbivore counter-adaptations to plant structural defence and illustrate how herbivore attack can induce the deployment of intensified defensive measures. Although there have been few studies detailing how plant defences vary with age, we show that allocation to structural defences is related to plant ontogeny. Age-related changes in the deployment of structural defences plus a paucity of appropriate studies are two reasons why relationships with other plant fitness characteristics may be obscured, although we describe studies where trade-offs between structural defence and plant growth, reproduction, and chemical defences have been demonstrated. We also show how resource availability influences the expression of structural defences and demonstrate how poorly our understanding of plant structural defence fits into contemporary plant defence theory. Finally, we suggest how a better understanding of plant structural defence, particularly within the context of plant defence syndromes, would not only improve our understanding of plant defence theory, but enable us to predict how plant morphological responses to climate change might influence interactions at the individual (plant growth trade-offs), species (competition), and ecosystem (pollination and herbivory) levels.     

植物邻体间的正相互作用

植物间的正负相互作用是构建植被群落的重要因素,也是群落生态学研究的中心内容之一。近20a来,植物间正相互作用的研究得到快速发展。综述了正相互作用的定义,不同植物群落中的直接、间接正相互作用及其发生机制,正相互作用研究的实验和模型方法,正负相互作用随胁迫梯度的变化及正相互作用对群落构建的影响。探讨了正相互作用研究前景:(1)进一步理解正负相互作用的平衡及其对群落构建的影响;(2)加深对全球变暖背景下的正相互作用的认识;(3)需把正相互作用研究同进化联系起来;(4)充分发挥正相互作用在生态系统中的推动力作用,把正相互作用应用到生态恢复中,为恢复退化生态系统服务。     

Gall insects and selection on plant vigor: can susceptibility compromise success in competition?

Gall insects select vigorously growing plants and plant parts when initiating gall formation. Vigor is associated with rapid growth rate, and in turn, rapid growth confers competitiveness. Are there conditions under which the cost of vigor, in the form of increased susceptibility to attack, outweighs the benefit of competitive success? I present a simulation model to explore the interaction between susceptibility and competition on the selective advantage of increased growth rate. Assuming size-symmetric competition, the model shows that in general, vigor is favored (benefit > cost) at low to intermediate gall loads. At very high plant densities, however, plants with high gall loads may lose standing in the competitive size hierarchy from which they cannot recover. The details of this result, however, change somewhat when competition is size-asymmetric, that is, when a larger focal plant suppresses smaller neighbors, but smaller neighbors cannot exert a reciprocal effect on the focal. At low densities, the pattern of selection on growth rate is qualitatively similar to the size-symmetric case. At higher plant densities, however, fast-growing genotypes can suppress slow ones so much during the preattack phase that even at the highest gall loads they maintain their standing in the competitive hierarchy. Thus, heavy gall insect attack on vigorous plants can impose selection against high intrinsic growth rates under strong symmetric competition, but not strong asymmetric competition. While life history traits can evolve as a correlated response to selection on defensive traits that reduce susceptibility, this model reveals that susceptibility can evolve as a correlated response to selection on basic life history traits.     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.