Effect of soil nutrient heterogeneity on the symmetry of belowground competition

Size variability in plant populations has been extensively studied andmuch of this inquiry has focused on the types of competition that are involvedin increasing or decreasing size inequality. It is often assumed thatcompetition for light is size-asymmetric, meaning that a plant canpotentially dominate a competitive relationship through shading if it is tallerthan its competitor. The light resources obtained by the taller plant are thusdisproportionate to its size. In contrast, competition for soil resources maybemore size-symmetric, the amount of soil nutrients obtained seems to be indirect proportion to a plant's size. Most studies examining belowgroundcompetition have used homogeneouslydistributed nutrient resources. Soil homogeneity could makesize-asymmetric belowground competition unlikely, but homogeneity is notoften found in nature. In this study I use a greenhouse pot experimentutilizingIpomoea tricolor to examine the hypothesis thatsize-asymmetric competition for soil nutrients may result when soilresource distribution is spatially heterogeneous. The results did not supportthe hypothesis of belowgroundsize-asymmetric competition. Differences between experimental treatmentsand controls were not statistically significant suggesting size-symmetriccompetition. Implications of these results are discussed.     

Soil phosphorus heterogeneity and mycorrhizal symbiosis regulate plant intra-specific competition and size distribution

We investigated the interactive effects of soil phosphorus (P) heterogeneity, plant density and mycorrhizal symbiosis on plant growth and size variability of Trifolium subterraneum. We set up mesocosms (trays 49Ꮉ cm and 12 cm deep) with the same amount of available P, but distributed either homogeneously or heterogeneously, in randomly arranged cells (7ǻ cm each) with high or low available P. The trays were planted with either 1 or 4 seedlings of T. subterraneum per cell. Half of the trays were inoculated with spores of the mycorrhizal fungus Gigaspora margarita. We harvested the plants when leaves just started to overlap, 8 weeks after planting. Plants growing in high P cells had the lowest percentage infection, but the highest mean shoot and root biomass and root length. The mean size of the plants in each cell was determined mainly by local P concentration. However, in plants growing in high density, low P cells, ca. 20% of the variability in plant biomass was explained by the number of adjacent cells with high P. Patchy trays had the highest total shoot biomass, independently of mycorrhizal infection or plant density. Inoculated trays (M) had higher total shoot biomass and relative competition intensity (measured as reduction in plant biomass due to increased density) than non-inoculated trays (NM). Plant density reduced the plant response to mycorrhizal infection, and its effect was independent of P distribution. All populations growing in patchy trays, and low density mycorrhizal ones, had the highest plant-size inequality, presumably because patchy distribution of P and mycorrhizal infection increased competitive asymmetry. We conclude that mycorrhizal symbiosis has the potential to strongly influence plant population structure when soil nutrient distribution is heterogeneous because it promotes pre-emption of limiting resources.     

Genetic variation for root architecture, nutrient uptake and mycorrhizal colonisation in Medicago truncatula accessions

Sustainable agriculture strives for healthy, high yielding plants with minimal agronomic inputs. Genetic solutions to increase nutrient uptake are desirable because they provide ongoing improvements. To achieve this it is necessary to identify genes involved in uptake and translocation of nutrients. We selected Medicago truncatula L. as a model because of its: i) close genetic relationship to food legumes, ii) use as a pasture legume in southern Australia and iii) availability of mapping populations generated from genetically diverse accessions. We discovered statistically significant differences between eight accessions for: root architecture in growth pouches, % root colonisation with the arbuscular mycorrhizal (AM) fungus Glomus intraradices, and plant tissue concentration of most macro- and micronutrients. Mycorrhizal colonisation had a significant effect on P concentration in roots but not shoots, Mg concentration in both roots and shoots, and the concentration of various micronutrients in shoots including Fe, Ca, but not Zn. Comparison of micronutrient uptake between root and shoot tissues showed that some M. truncatula accessions were more efficient at mobilisation of nutrients from roots to shoots. We are now in a position to use existing mapping populations of M. truncatula to identify quantitative trait loci important for human health and sustainable agriculture.     

Effects of soil nutrient heterogeneity on intraspecific competition in the invasive, clonal plant Alternanthera philoxeroides

Background and Aims

Fine-scale, spatial heterogeneity in soil nutrient availability can increase the growth of individual plants, the productivity of plant communities and interspecific competition. If this is due to the ability of plants to concentrate their roots where nutrient levels are high, then nutrient heterogeneity should have little effect on intraspecific competition, especially when there are no genotypic differences between individuals in root plasticity. We tested this hypothesis in a widespread, clonal species in which individual plants are known to respond to nutrient heterogeneity.

Methods

Plants derived from a single clone of Alternanthera philoxeroides were grown in the greenhouse at low or high density (four or 16 plants per 27·5 × 27·5-cm container) with homogeneous or heterogeneous availability of soil nutrients, keeping total nutrient availability per container constant. After 9 weeks, measurements of size, dry mass and morphology were taken.

Key Results

Plants grew more in the heterogeneous than in the homogeneous treatment, showing that heterogeneity promoted performance; they grew less in the high- than in the low-density treatment, showing that plants competed. There was no interactive effect of nutrient heterogeneity and plant density, supporting the hypothesis that heterogeneity does not affect intraspecific competition in the absence of genotypic differences in plasticity. Treatments did not affect morphological characteristics such as specific leaf area or root/shoot ratio.

Conclusions

Results indicate that fine-scale, spatial heterogeneity in the availability of soil nutrients does not increase competition when plants are genetically identical, consistent with the suggestion that effects of heterogeneity on competition depend upon differences in plasticity between individuals. Heterogeneity is only likely to increase the spread of monoclonal, invasive populations such as that of A. philoxeroides in China.     

Belowground interactions between Kalmia angustifolia and Picea mariana: roles of competition,root exudates and ectomycorrhizal association

Plant and Soil - Belowground competition and allelopathic interference of neighbouring plants play important roles in shaping plant performance. We assessed the effect of belowground interactions...     

Plant root response to heterogeneity of soil resources: Effects of nutrient patches, AM symbiosis, and species composition

Response of the underground parts of a whole grassland community to heterogeneity in soil resources was studied under field conditions. The increased availability of nutrients (N, P) in patches was combined with suppression of arbuscular mycorrhizal symbiosis (by fungicide benomyl) and with modification of substrate structure (sieved soil, sieved soil mixed with coarse or fine sand). Three experimental runs were established in 1999, each with seven replicate blocks. Root and rhizome biomass was determined before the experimental treatment and after 53 days of the experiment. Plant community composition in neighbourhood of patches was recorded. Nutrient application had the most pronounced effect on the root biomass grown into the patches (the root biomass increased 1.64 times). The root biomass in patches with fungicide application was 1.17 times higher than in the untreated patches. The modification of substrate structure had no effect on the root biomass in patches but it did modify the effect of benomyl. The amount of root biomass proliferating into the patches was not related to species composition of vegetation near the patches. However, higher frequency of grass tillers in the patch neighbourhood and the mycorrhizal status of neighbouring species correlated significantly with the amount of root biomass in the experimental patches. In the pre-treatment community, amount of root biomass was significantly related to the abundance of grasses and mycorrhizal forbs surrounding the sampling points.     

Simulation of nutrient uptake by a growing root system considering increasing root density and inter-root competition

A simulation model is presented which describes uptake of a growth limiting nutrient from soil by a growing root system. The root surface is supposed to behave like a zero-sink. Uptake of the nutrient is therefore determined by the rate of nutrient supply to the root surface by mass flow and diffusion. Inter-root competition and time dependent root density are accounted for by assigning to each root a finite cylindrical soil volume that delivers nutrients. The radius of these cylinders declines with increasing root density. Experiments with rape plants grown on quartz sand were used to evaluate the model. Simulated nitrogen uptake agreed well with observed uptake under nitrogen limiting conditions. In case no nitrogen limitation occurred nitrogen uptake was overestimated by the model, probably because the roots did not behave like a zero-sink any more.     

Functional aspects of root architecture and mycorrhizal inoculation with respect to nutrient uptake capacity

The aim of this research was to investigate the effect of arbuscular mycorrhizal (AM) colonisation on root morphology and nitrogen uptake capacity of carob ( Ceratonia siliqua L.) under high and low nutrient conditions. The experimental design was a factorial arrangement of presence/absence of mycorrhizal fungus inoculation ( Glomus intraradices) and high/low nutrient status. Percent AM colonisation, nitrate and ammonium uptake capacity, and nitrogen and phosphorus contents were determined in 3-month-old seedlings. Grayscale and colour images were used to study root morphology and topology, and to assess the relation between root pigmentation and physiological activities. AM colonisation lead to a higher allocation of biomass to white and yellow parts of the root. Inorganic nitrogen uptake capacity per unit root length and nitrogen content were greatest in AM colonised plants grown under low nutrient conditions. A better match was found between plant nitrogen content and biomass accumulation, than between plant phosphorus content and biomass accumulation. It is suggested that the increase in nutrient uptake capacity of AM colonised roots is dependent both on changes in root morphology and physiological uptake potential. This study contributes to an understanding of the role of AM fungi and root morphology in plant nutrient uptake and shows that AM colonisation improves the nitrogen nutrition of plants, mainly when growing at low levels of nutrients.     

Selenium as an anticancer nutrient: roles in cell proliferation and tumor cell invasion

Selenium is an essential dietary component for animals including humans, and there is increasing evidence for the efficacy of certain forms of selenium as cancer-chemopreventive compounds. In addition, selenium appears to have a protective effect at various stages of carcinogenesis including both the early and later stages of cancer progression. Mechanisms for selenium-anticancer action are not fully understood; however, several have been proposed: antioxidant protection, enhanced carcinogen detoxification, enhanced immune surveillance, modulation of cell proliferation (cell cycle and apoptosis), inhibition of tumor cell invasion and inhibition of angiogenesis. Research has shown that the effectiveness of selenium compounds as chemopreventive agents in vivo correlates with their abilities to affect the regulation of the cell cycle, to stimulate apoptosis and to inhibit tumor cell migration and invasion in vitro. This article reviews the status of knowledge concerning selenium metabolism and its anticancer effects with particular reference to the modulation of cell proliferation and the inhibition of tumor cell invasion.     

Root proliferation and seed yield in response to spatial heterogeneity of below-ground competition

Here, we tested the predictions of a 'tragedy of the commons' model of below-ground plant competition in annual plants that experience spatial heterogeneity in their competitive environment. Under interplant competition, the model predicts that a plant should over-proliferate roots relative to what would maximize the collective yield of the plants. We predict that a plant will tailor its root proliferation to local patch conditions, restraining root production when alone and over-proliferating in the presence of other plants. A series of experiments were conducted using pairs of pea (Pisum sativum) plants occupying two or three pots in which the presence or absence of interplant root competition was varied while nutrient availability per plant was held constant. In two-pot experiments, competing plants produced more root mass and less pod mass per individual than plants grown in isolation. In three-pot experiments, peas modulated this response to conditions at the scale of individual pots. Root proliferation in the shared pot was higher compared with the exclusively occupied pot. Plants appear to display sophisticated nutrient foraging with outcomes that permit insights into interplant competition.     

Lateral root function and root overlap among mycorrhizal and nonmycorrhizal herbs in a Florida shrubland, measured using rubidium as a nutrient analog

Lateral root spread is an important indicator of the area over which plants forage belowground and has implications for individual plant fitness, belowground competition, and ultimately both population dynamics and community structure. Because fine lateral roots are difficult to excavate and measure, we used rubidium (Rb), an analog for potassium that occurs at very low levels in soils and plants, as a tracer of root activity. We injected Rb into sandy soils of a xeric Florida shrubland and examined uptake by four small herbaceous species that generally do not overlap aboveground. All four species took up Rb at distances of up to 97 cm (with an average lateral root spread of 51 cm), indicating that belowground neighborhoods are larger than aboveground. The arbuscular mycorrhizal species exhibited greater Rb enrichment without any change in lateral spread. The spatial arrangement of plants with elevated levels of Rb relative to Rb patches was consistent with root and mycorrhizal systems that are radially discontinuous and asymmetric. Furthermore, many individuals acquired Rb from the same patches, suggesting overlap in root function and the potential for belowground competition, although small-scale spatial and/or temporal segregation could have occurred.     

Interactions between dispersal, competition, and landscape heterogeneity

It is widely acknowledged that space has an important role in population regulation, yet more specific knowledge into how the relevant factors interact attains little consensus. We address this issue via a stochastic, individual based model of population dynamics, in a continuous space continuous time framework. We represent habitat quality as a continuously varying surface over the two-dimensional landscape, and assume that the quality affects either fecundity (rate of propagule production) or probability of propagule establishment. We control the properties of the landscape by two parameters, which we call the patch size (the characteristic length scale in quality variation), and the level of heterogeneity (the characteristic quality difference between poor quality and high quality areas). In addition to such exogenous variability, we also account for endogenous factors causing spatial variation by assuming localised dispersal and competition. We find that heterogeneity has a general positive effect on population density, and hence it is beneficial to improve best quality habitat at the expense of worst quality habitat. With regards to patch size, we find an intermediate optimum, due to a conflict between minimising the loss of propagules to low quality regions and maximising the benefits of heterogeneity. We address the consequences of regional stochasticity by allowing the environmental conditions change in time. The cost of having to continuously track where the favourable conditions have moved to ultimately reduces population size.     

Rheumatoid synovium: its proliferation, heterogeneity and polarization

All of the changes observed suggest that a viral agent may form the basis for these alterations [in rheumatoid synovium]. However, studies to date have failed to isolate or identify the putative causative virus and the search continues.     

Meeting nutrient requirements: the roles of power and efficiency

Conclusions We have attempted to illustrate the importance of nutrient transfer functions and their key quantitative parameters, power and efficiency. While this brief account has focused on one aspect of nutritional behaviour, the duration of intermeal intervals, the models are equally useful for understanding other components of feeding behaviour such as meal size and food switching. Like-wise, they provide a framework for formulating quantitative predictions about the dynamics of the physiological processes involved in digestion, absorption and the utilisation of food. These aspects are dealt with more fully elsewhere.     

An arbuscular mycorrhizal inoculum enhances root proliferation in, but not nitrogen capture from, nutrient-rich patches in soil

Most work on root proliferation to a localized nutrient supply has ignored the possible role of mycorrhizal fungi, despite their key role in nutrient acquisition. Interactions between roots of Plantago lanceolata , an added arbuscular mycorrhiza (AM) inoculum and nitrogen capture from an organic patch ( Lolium perenne shoot material) dual-labelled with ¹⁵N and ¹³C were investigated, to determine whether root proliferation and nitrogen (N) capture was affected by the presence of AM fungi. Decomposition of the organic patch in the presence and absence of roots peaked in all treatments at day 3, as shown by the amounts of ¹³CO₂ detected in the soil atmosphere. Plant N concentrations were higher in the treatments with added inoculum 10 d after patch addition, but thereafter did not differ among treatments. Plant phosphorus concentrations at the end of the experiment were depressed by the addition of the organic residue in the absence of mycorrhizal inoculum. Although uninoculated plants were also colonized by mycorrhizal fungi, colonization was enhanced at all times by the added inoculum. Addition of the AM inoculum increased root production, observed in situ by the use of minirhizotron tubes, most pronouncedly within the organic patch zone. Patch N capture by the end of the experiment was c . 7.5% and was not significantly different as a result of adding an AM inoculum. Furthermore, no ¹³C enrichments were detected in the plant material in any of the treatments showing that intact organic compounds were not taken up. Thus, although the added AM fungal inoculum benefited P. lanceolata seedlings in terms of P concentrations of tissues it did not increase total N capture or affect the form in which N was captured by P. lanceolata roots.     

Predation, competition, and nutrient recycling: a stoichiometric approach with multiple nutrients

A model for two competing prey species and one predator is formulated in which three essential nutrients can limit growth of all populations. Prey take up dissolved nutrients and predators ingest prey, assimilating a portion of ingested nutrients and recycling or respiring the balance. For all species, the nutrient contents of individuals vary and growth is coupled to increasing content of the limiting nutrient. This model was parameterized to describe a flagellate preying on two bacterial species, with carbon (C), nitrogen (N), and phosphorus (P) as nutrients. Parameters were chosen so that the two prey species would stably coexist without predators under some nutrient supply conditions. Using numerical simulations, the long-term outcomes of competition and predation were explored for a gradient of N:P supply ratios, varying C supply, and varying preference of the predator for the two prey. Coexistence and competitive exclusion both occurred under some conditions of nutrient supply and predator preference. As in simpler models of competition and predation these outcomes were largely governed by apparent competition mediated by the predator, and resource competition for nutrients whose effective supply was partly governed by nutrient recycling also mediated by the predator. For relatively small regions of parameter space, more complex outcomes with multiple attractors or three-species limit cycles occurred. The multiple constraints posed by multiple nutrients held the amplitudes of these cycles in check, limiting the influence of complex dynamics on competitive outcomes for the parameter ranges explored.     

Effects of a vesicular-arbuscular mycorrhizal fungus and other soil microorganisms on growth,mineral nutrient acquisition and root exudation of soil-grown maize plants

Maize (Zea mays L. cv. Alize) plants were grown in a calcareous soil in pots divided by 30-m nylon nets into three compartments, the central one for root growth and the outer ones for hyphal growth. Sterle soil was inoculated with either (1) rhizosphere microorganisms other than vesicular-arbuscular mycorrhizal (VAM) fungi, (2) rhizosphere microorganisms together with a VAM fungus [Glomus mosseae (Nicol. and Gerd.) Gerdemann and Trappel], or (3) with a gamma-irradiated inoculum as control. Plants were grown under controlled-climate conditions and harvested after 3 or 6 weeks. VAM plants had higher shootroot ratios than non-VAM plants. After 6 weeks, the concentrations of P, Zn and Cu in roots and shoots had significantly increased with VAM colonization, whereas Mn concentrations had significantly decreased. Root exudates were collected on agar sheets placed on the interface between root and hyphal compartments. Six-week-old VAM and non-VAM plants had similar root exudate compositions of 72–73% reducing sugars, 17–18% phenolics, 7% organic acids and 3% amino acids. In another experiment in which root exudates were collected on agar sheets with or without antibiotics, the amounts of amino acids and carbohydrates recovered were similar in VAM and non-VAM plants. However, threeto sixfold higher amounts of carbohydrates, amino acids and phenolics were recovered when antibiotics were added to the agar sheets. Thus, the high microbial activity in the rhizosphere and on the rhizoplane limits the exudates recovered from roots.     

Phosphate in the arbuscular mycorrhizal symbiosis: transport properties and regulatory roles

In response to the colonization by arbuscular mycorrhizal (AM) fungi, plants reprioritize their phosphate (Pi)-uptake strategies to take advantage of nutrient transfer via the fungus. The mechanisms underlying Pi transport are beginning to be understood, and recently, details of the regulation of plant and fungal Pi transporters in the AM symbiosis have been revealed. This review summarizes recent advances in this area and explores current data and hypotheses of how the plant Pi status affects the symbiosis. Finally, suggestions of an interrelationship of Pi and nitrogen (N) in the AM symbiosis are discussed.     

Effects of nutrient supply and competition from other species on root growth of Lolium perenne in soil

Summary In competition experiments with Lolium perenne and Agrostis tenuis on sandy soil with nitrogen supplied and therefore not limiting, it was found that the competitive interactions could be explained in terms of phosphate uptake, and that the ratio of root weight: length was proportional to root density. The effects of competition were then investigated in an experimental system that enabled them to be distinguished from those of nutrient supply. High levels of nutrients specifically stimulated the production of fine laterals whereas competition affected length and weight of the root system of Lolium equally. The ecological implications are discussed in the light of recent physiological work on root responses to nutrient supply. re]19750324