A simulation study of the effects of architectural constraints and resource translocation on population structure and competition in clonal plants

(1) Spatially explicit simulation of clonal plant growth is used to determine how ramet-level traits affect ramet density, spatial pattern of ramets and competitive ability of a clonal plant. The simulation model used combines elements of (i) an individual-based model of plant interactions, (ii) an architectural model of clonal plant growth, and (iii) a model of resource translocation within a set of physiologically integrated plant individuals. (2) The effects of two groups of parameters were studied: growth and resource acquisition parameters (resource accumulation, density-dependence of resource accumulation, resource translocation between ramets) and architectural rules (branching angle and probability of branching, internode length). The model was parameterised by values approximating those of clonally growing grasses as closely as possible. The basic parameter values were chosen from a short-turf grassland. Sensitivity analysis was carried out on relevant parameters around three basic points in the parameter space. Both single-species and two-species systems were studied. (3) It is shown that increasing resource acquisition and growth parameters increase ramet density, genet number and competitive ability. Translocation parameters and architectural parameters modify the effects of resource acquisition and growth, but their effect in single-species stands was smaller. (4) The simulations of species with fixed ramet sizes showed that ramet density in single-species stands cannot be used for predicting competitive ability. Increase in resource acquisition and growth parameters was correlated with an increase in equilibrium ramet density and competitive ability. Increasing branching angle, branching probability or internode length lead to an increased competitive ability, but did not affect equilibrium ramet density. Change of architectural parameters could therefore affect competitive ability independently of their effect on the final ramet density. (5) Spatial pattern both in single-species and two-species stands was also highly parameter-dependent. Changes in architectural parameters and in translocation usually lead to pronounced change in the spatial pattern; change in growth and resource acquisition parameters generally had little effect on spatial pattern.     

A dynamic model of plant growth with interactions between development and functional mechanisms to study plant structural plasticity related to trophic competition

Background and Aims

The strong influence of environment and functioning on plant organogenesis has been well documented by botanists but is poorly reproduced in most functional–structural models. In this context, a model of interactions is proposed between plant organogenesis and plant functional mechanisms.

Methods

The GreenLab model derived from AMAP models was used. Organogenetic rules give the plant architecture, which defines an interconnected network of organs. The plant is considered as a collection of interacting ‘sinks’ that compete for the allocation of photosynthates coming from ‘sources’. A single variable characteristic of the balance between sources and sinks during plant growth controls different events in plant development, such as the number of branches or the fruit load.

Key Results

Variations in the environmental parameters related to light and density induce changes in plant morphogenesis. Architecture appears as the dynamic result of this balance, and plant plasticity expresses itself very simply at different levels: appearance of branches and reiteration, number of organs, fructification and adaptation of ecophysiological characteristics.

Conclusions

The modelling framework serves as a tool for theoretical botany to explore the emergence of specific morphological and architectural patterns and can help to understand plant phenotypic plasticity and its strategy in response to environmental changes.Key words: Trophic plasticity, plant growth, functional–structural models, dynamic system, interactions, GreenLab     

Size-asymmetric competition and size-asymmetric growth in a spatially explicit zone-of-influence model of plant competition

Size-asymmetric competition among plants is usually defined as resource pre-emption by larger individuals, but it is usually observed and measured as a disproportionate size advantage in the growth of larger individuals in crowded populations (“size-asymmetric growth”). We investigated the relationship between size-asymmetric competition and size-asymmetric growth in a spatially explicit, individual-based plant competition model based on overlapping zones of influence (ZOI). The ZOI of each plant is modeled as a circle, growing in two dimensions. The size asymmetry of competition is reflected in the rules for dividing up the overlapping areas. We grew simulated populations with different degrees of size-asymmetric competition and at different densities and analyzed the size dependency of individual growth by fitting coupled growth functions to individuals. The relationship between size and growth within the populations was summarized with a parameter that measures the size asymmetry of growth. Complete competitive symmetry (equal division of contested resources) at the local level results in a very slight size asymmetry in growth. This slight size asymmetry of growth did not increase with increasing density. Increased density resulted in increased growth asymmetry when resource competition at the local level was size asymmetric to any degree. Size-asymmetric growth can be strong evidence that competitive mechanisms are at least partially size asymmetric, but the degree of size-asymmetric growth is influenced by the intensity as well as the mode of competition. Intuitive concepts of size-asymmetric competition among individuals in spatial and nonspatial contexts are very different.     

病原体感染对物种间资源竞争的影响——基于资源竞争理论

病原体感染对种间竞争的影响可能是因为改变了宿主的资源利用过程,然而竞争模型（Lotka-Volterra）由于参数化竞争系数而忽略了资源的动态变化过程,因此基于此类模型的研究无法揭示病原体对宿主资源利用的影响。基于Tilman的资源竞争理论构建了病原体感染一个物种的资源竞争模型,通过分析宿主物种资源利用效率的变化探讨了病原体对种间竞争的影响。结果表明：（1）病原体降低了宿主对资源的消耗率（消费矢量变短）,抬高了对资源的最低需求（零等倾线上移）,这意味着宿主的竞争力减弱;（2）虽然感染影响了竞争物种的密度,但不会改变共存物种的共存状态;（3）病原体可以使宿主物种的竞争对手更容易入侵,形成共存局面,极大地扩大了竞争物种共存的参数范围,本质上促进了物种多样性维持;（4）病原体的传播率和毒性也复杂地影响了竞争物种共存,传播率越大越能促进物种共存,而中等强度毒性最能促进物种共存。研究结果明确了病原体对物种资源利用模式的潜在改变,强调了病原体在物种共存和生物多样性维持中的重要性。     

Plant population growth and competition in a light gradient: a mathematical model of canopy partitioning

Can a difference in the heights at which plants place their leaves, a pattern we call canopy partitioning, make it possible for two competing plant species to coexist? To find out, we examine a model of clonal plants living in a nonseasonal environment that relates the dynamical behavior and competitive abilities of plant populations to the structural and functional features of the plants that form them. This examination emphasizes whole plant performance in the vertical light gradient caused by self-shading. This first of three related papers formulates a prototype single species Canopy Structure Model from biological first principles and shows how all plant properties work together to determine population persistence and equilibrium abundance. Population persistence is favored, and equilibrium abundance is increased, by high irradiance, high maximum photosynthesis rate, rapid saturation of the photosynthetic response to increased irradiance, low tissue respiration rate, small amounts of stem and root tissue necessary to support the needs of leaves, and low density of leaf, stem, and root tissues. In particular, equilibrium abundance decreases as mean leaf height increases because of the increased cost of manufacturing and maintaining stem tissue. All conclusions arise from this formulation by straightforward analysis. The argument concludes by stating this formulation's straightforward extension, called a Canopy Partitioning Model, to two competing species.     

Biophysical aspects of resource acquisition and competition in algal mixotrophs

Mixotrophic organisms combine autotrophic and heterotrophic nutrition and are abundant in both freshwater and marine environments. Recent observations indicate that mixotrophs constitute a large fraction of the biomass, bacterivory, and primary production in oligotrophic environments. While mixotrophy allows greater flexibility in terms of resource acquisition, any advantage must be traded off against an associated increase in metabolic costs, which appear to make mixotrophs uncompetitive relative to obligate autotrophs and heterotrophs. Using an idealized model of cell physiology and community competition, we identify one mechanism by which mixotrophs can effectively outcompete specialists for nutrient elements. At low resource concentrations, when the uptake of nutrients is limited by diffusion toward the cell, the investment in cell membrane transporters can be minimized. In this situation, mixotrophs can acquire limiting elements in both organic and inorganic forms, outcompeting their specialist competitors that can utilize only one of these forms. This advantage can be enough to offset as much as a twofold increase in additional metabolic costs incurred by mixotrophs. This mechanism is particularly relevant for the maintenance of mixotrophic populations and productivity in the highly oligotrophic subtropical oceans.     

Herbivory and plant resource competition: a review of two interacting interactions

This review discusses the prevalence and potential for interactive effects between herbivory and competition on plant growth and biomass, and it is apparent that such effects typically arise when there is a mismatch between the spatial scale of herbivore behaviour (food or patch choice) and the spatial heterogeneity of the plant community. Historically, such interactive effects have been examined using two approaches. Studies using the first approach have excluded plant neighbors and herbivores in a factorial experiment, and scored effects on plant biomass. Studies using the second approach have observed herbivore abundance or herbivory on plants with or without plant neighbors, and have identified a large number of mechanisms underlying such interactive effects. The two types of studies have produced somewhat conflicting results, where interactive effects have been commonly observed in studies using the second approach and only rarely in studies using the first approach. This is most likely a consequence of a biased choice of study systems, where studies using the first approach have primarily studied mammalian herbivory while studies using the second approach have been more focussed on insect herbivory. Moreover, studies using the first approach have typically been very small-scale manipulations and this probably precludes most possible interactive effects in systems with mammalian herbivory. This points to the fact that studies examining interactive effects of herbivory and plant competition should more carefully consider the behaviour and life history of herbivores included in the study prior to the design of removal experiments.     

Relationships among three pathways for resource acquisition and their contribution to plant performance in the emergent aquatic Plant Lythrum salicaria (L.)

Three pathways for resource acquisition exist in the emergent aquatic plant, Lythrum salicaria (L.); a subterranean root system, a free-floating adventitious root system, and arbuscular mycorrhiza (AM) fungal hyphae colonizing subterranean roots. This study examined the relationship(s) among these pathways and their contribution to plant performance. If the free-floating adventitious root system and/or AM fungi contribute to plant growth in wetland habitats, we predicted that their absence would result in a significant reduction in plant performance. Furthermore, if a reduction in resource uptake, effected by an absence of free-floating adventitious roots and/or AM fungi, is compensated for by increased allocation to remaining pathway(s) for resource uptake, we predicted altered patterns of resource allocation among shoots and the remaining pathway(s) for resource uptake. Contrary to our predications, plants experiencing adventitious root removal and/or grown in the absence of AM fungi generally had greater biomass and total shoot height than controls. Similarly, while levels of AM colonization and subterranean root biomass displayed a treatment effect, the observed responses did not correspond with our predictions. This was also true for shoot : subterranean root dry weight ratios. Our results indicate that there is interaction among the 3 pathways for resource acquisition in L. salicaria and an effect on plant performance. The adaptive significance of these characteristics is unclear, highlighting the potential difficulties in extrapolating from terrestrial to aquatic plant species and among aquatic plant species with potentially different life history strategies.     

微重力环境影响植物生长发育的研究进展

微重力是最独特的空间环境条件之一,研究微重力对不同植物种类以及不同植物部位的影响是空间生物学的重要内容之一,对于建立生物再生式生命保障系统意义重大。生物再生式生命保障系统是未来开展长期载人空间活动的核心技术,其优势在于能在一个密闭的系统内持续再生氧气,水和食物等高等动物生活必需品,植物部件是生物再生式生命保障系统的重要组成部分。了解和掌握微重力对植物生长发育的影响,有助于采取有效的作业制度确保其正常生长发育和繁殖,是成功建立生物再生式生命保障系统的首要关键。该文就植物在空间探索中的地位和作用,地面模拟微重力的装置以及国内外有关微重力对植物的影响做一综述。现有的研究结果包括,未来长期的载人航天任务需要植物通过光合作用为生物再生式生命保障系统提供部分动物营养、洁净水以及清除系统中的固体废物和二氧化碳;三维随机回旋装置是目前地面上模拟微重力效应的主要装置之一,尤其适用于植物材料的长期模拟微重力处理;国内外有关微重力对植物影响的报道生理生化水平多集中在植物的生长发育和生理反应,比如表型变化或者与重力相关的激素或者钙离子的再分配,细胞或亚细胞水平主要有细胞壁、线粒体、叶绿体以及细胞骨架等,基因和蛋白质表达水平的研究对象主要为拟南芥。由于实验方法和材料之间的差异,微重力对不同植物或者植物不同部位在各个水平的影响效果并不一致,未来需要开展更多的相关研究工作。     

Shedding light on plant competition: modelling the influence of plant morphology on light capture (and vice versa)

A plant's morphology is both strongly influenced by local light availability and, simultaneously, strongly influences this local light availability. This reciprocal relationship is complex, but lies at the heart of understanding plant growth and competition. Here, we develop a sub-individual-based simulation model, cast at the level of interacting plant components. The model explicitly simulates growth, development and competition for light at the level of leaves, branches, etc., located in 3D space. In this way, we are able to explore the manner in which the low-level processes governing plant growth and development give rise to individual-, cohort-, and community-level phenomena. In particular, we show that individual-level trade-offs between growing up and growing out arise naturally in the model, and robustly give rise to cohort-level phenomena such as self-thinning, and community processes such as the effect of ecological disturbance on the maintenance of biodiversity. We conclude with a note on our methodology and how to interpret the results of simulation models such as this one.     

Earthworm and belowground competition effects on plant productivity in a plant diversity gradient

Diversity is one major factor driving plant productivity in temperate grasslands. Although decomposers like earthworms are known to affect plant productivity, interacting effects of plant diversity and earthworms on plant productivity have been neglected in field studies. We investigated in the field the effects of earthworms on plant productivity, their interaction with plant species and functional group richness, and their effects on belowground plant competition. In the framework of the Jena Experiment we determined plant community productivity (in 2004 and 2007) and performance of two phytometer plant species [Centaurea jacea (herb) and Lolium perenne (grass); in 2007 and 2008] in a plant species (from one to 16) and functional group richness gradient (from one to four). We sampled earthworm subplots and subplots with decreased earthworm density and reduced aboveground competition of phytometer plants by removing the shoot biomass of the resident plant community. Earthworms increased total plant community productivity (+11%), legume shoot biomass (+35%) and shoot biomass of the phytometer C. jacea (+21%). Further, phytometer performance decreased, i.e. belowground competition increased, with increasing plant species and functional group richness. Although single plant functional groups benefited from higher earthworm numbers, the effects did not vary with plant species and functional group richness. The present study indicates that earthworms indeed affect the productivity of semi-natural grasslands irrespective of the diversity of the plant community. Belowground competition increased with increasing plant species diversity. However, belowground competition was modified by earthworms as reflected by increased productivity of the phytometer C. jacea. Moreover, particularly legumes benefited from earthworm presence. Considering also previous studies, we suggest that earthworms and legumes form a loose mutualistic relationship affecting essential ecosystem functions in temperate grasslands, in particular decomposition and plant productivity. Further, earthworms likely alter competitive interactions among plants and the structure of plant communities by beneficially affecting certain plant functional groups.     

Nitrogen acquisition by roots: physiological and developmental mechanisms ensuring plant adaptation to a fluctuating resource

Background

Nitrogen (N) is one of the key mineral nutrients for plants and its availability has a major impact on their growth and development. Most often N resources are limiting and plants have evolved various strategies to modulate their root uptake capacity to compensate for both spatial and temporal changes in N availability in soil. The main N sources for terrestrial plants in soils of temperate regions are in decreasing order of abundance, nitrate, ammonium and amino acids. N uptake systems combine, for these different N forms, high- and low-affinity transporters belonging to multige families. Expression and activity of most uptake systems are regulated locally by the concentration of their substrate, and by a systemic feedback control exerted by whole-plant signals of N status, giving rise to a complex combinatory network. Besides modulation of the capacity of transport systems, plants are also able to modulate their growth and development to maintain N homeostasis. In particular, root system architecture is highly plastic and its changes can greatly impact N acquisition from soil.

Scope

In this review, we aim at detailing recent advances in the identification of molecular mechanisms responsible for physiological and developmental responses of root N acquisition to changes in N availability. These mechanisms are now unravelled at an increasing rate, especially in the model plant Arabidopsis thaliana L.. Within the past decade, most root membrane transport proteins that determine N acquisition have been identified. More recently, molecular regulators in nitrate or ammonium sensing and signalling have been isolated, revealing common regulatory genes for transport system and root development, as well as a strong connection between N and hormone signalling pathways.

Conclusion

Deciphering the complexity of the regulatory networks that control N uptake, metabolism and plant development will help understanding adaptation of plants to sub-optimal N availability and fluctuating environments. It will also provide solutions for addressing the major issues of pollution and economical costs related to N fertilizer use that threaten agricultural and ecological sustainability.     

Quantifying the effects of ozone on plant reproductive growth and development

Tropospheric ozone (O₃) is a harmful air pollutant that can negatively impact plant growth and development. Current O₃ concentrations ([O₃]) decrease forest productivity and crop yields and future [O₃] will likely increase if current emission rates continue. However, the specific effects of elevated [O₃] on reproductive development, a critical stage in the plant's lifecycle, have not been quantitatively reviewed. Data from 128 peer‐reviewed articles published from 1968 to 2010 describing the effects of O₃ on reproductive growth and development were analysed using meta‐analytic techniques. Studies were categorized based on experimental conditions, photosynthetic type, lifecycle, growth habit and flowering class. Current ambient [O₃] significantly decreased seed number (?16%), fruit number (?9%) and fruit weight (?22%) compared to charcoal‐filtered air. In addition, pollen germination and tube growth were decreased by elevated [O₃] compared to charcoal‐filtered air. Relative to ambient air, fumigation with [O₃] between 70 and 100 ppb decreased yield by 27% and individual seed weight by 18%. Reproductive development of both C₃ and C₄ plants was sensitive to elevated [O₃], and lifecycle, flowering class and reproductive growth habit did not significantly affect a plant's response to elevated [O₃] for many components of reproductive development. However, elevated [O₃] decreased fruit weight and fruit number significantly in indeterminate plants, and had no effect on these parameters in determinate plants. While gaps in knowledge remain about the effects of O₃ on plants with different growth habits, reproductive strategies and photosynthetic types, the evidence strongly suggests that detrimental effects of O₃ on reproductive growth and development are compromising current crop yields and the fitness of native plant species.     

Intraspecific competition in a reef coral: effects on growth and reproduction

Summary Growth rates and reproduction of a branching coral (Stylophora pistillata) were compared in the presence and in the absence of intraspecific competition. Field experiments demonstrated a significant decline in the growth rate of competing colonies compared to noncompeting control colonies; the growth rate slowed in all of the interacting individuals, irrespective of their place in the hierarchy of the intraspecific dominance or of their color morph. In case of immediate killing of the subordinate, the dominant colony grew at a normal rate. In addition to the marked decrease in the growth rate of interacting colonies, the typical symmetry shape of these colonies was changed to an abnormal growth form. The number of female gonads per polyp was significantly reduced in colonies competing intraspecifically, and the typical synchrony in reproduction among different branches of a given colony was changed and desynchronized. Again, these results did not correlate with the hierarchy of dominance. We conclude that intraspecific competition in reef corals involves great investment of energy. The ecological significance and the different pathways of this competition are discussed.     

Intra- and intergenotypic competition in Drosophila melanogaster: effects of density on larval survival and rate of development

We have examined the effects of density and frequency in the larval competition of Drosophila melanogaster by measuring three fitness components: viability (V), mean development time (MDT) and a combination of these two (E). We have detected (contrary to most published results) non-linear effects of density in single-genotype cultures; in addition, different functions are required to describe the density effects below and above the optimal density. Frequency has also non-linear effects in the two-genotype cultures. Only one polymorphic equilibrium frequency, which is stable, occurs with respect to V; but two polymorphic equilibria, one stable and one unstable, exist with respect to E. The responses in single-genotype cultures do not allow one to predict the outcome of the competition in two-genotype cultures.     

Parameter stability of the functional-structural plant model GREENLAB as affected by variation within populations, among seasons and among growth stages

BACKGROUND AND AIMS: It is increasingly accepted that crop models, if they are to simulate genotype-specific behaviour accurately, should simulate the morphogenetic process generating plant architecture. A functional-structural plant model, GREENLAB, was previously presented and validated for maize. The model is based on a recursive mathematical process, with parameters whose values cannot be measured directly and need to be optimized statistically. This study aims at evaluating the stability of GREENLAB parameters in response to three types of phenotype variability: (1) among individuals from a common population; (2) among populations subjected to different environments (seasons); and (3) among different development stages of the same plants. METHODS: Five field experiments were conducted in the course of 4 years on irrigated fields near Beijing, China. Detailed observations were conducted throughout the seasons on the dimensions and fresh biomass of all above-ground plant organs for each metamer. Growth stage-specific target files were assembled from the data for GREENLAB parameter optimization. Optimization was conducted for specific developmental stages or the entire growth cycle, for individual plants (replicates), and for different seasons. Parameter stability was evaluated by comparing their CV with that of phenotype observation for the different sources of variability. A reduced data set was developed for easier model parameterization using one season, and validated for the four other seasons. KEY RESULTS AND CONCLUSIONS: The analysis of parameter stability among plants sharing the same environment and among populations grown in different environments indicated that the model explains some of the inter-seasonal variability of phenotype (parameters varied less than the phenotype itself), but not inter-plant variability (parameter and phenotype variability were similar). Parameter variability among developmental stages was small, indicating that parameter values were largely development-stage independent. The authors suggest that the high level of parameter stability observed in GREENLAB can be used to conduct comparisons among genotypes and, ultimately, genetic analyses.     

The effects of chrysomelid beetle grazing and plant competition on the growth of Rumex obtusifolius

Summary The interaction between grazing by Gastrophysa viridula and interspecific plant competition was investigated for Rumex obtusifolius growing in the field. During an eight-month growing season non-competing R. obtusifolius grew larger than competing plants although herbivore loads (beetle productivity per unit leaf area) were similar for competing and non-competing plants. However, grazing significantly reduced the growth of R. obtusifolius only in competing plants thus demonstrating a synergism between plant competition and invertebrate herbivory. The relevance of these findings for the distribution of plant species is discussed.     

Studies on plant growth-regulating substances.: The effects of light and hormone inhibitors on plant root growth

Light inhibition of primary root elongation in Zea mays seedlings was found to be an almost instantaneous response and the inhibitory effect was shown to persist for at least six hours following a short exposure to white light. Removal of the root cap completely removed the inhibitory effect of light. The apical region of the root appeared to be mainly responsible for perceiving light and initiating the production of factors involved in the inhibition of elongation. Extraction of the roots of light-exposed Zea seedlings revealed the presence of the carotenoid pigments, violaxanthin, neoxanthin and lutein, together with an unidentified non-acidic growth-inhibitory compound. Xanthoxin, however, was not detected.