Nutrient Heterogeneity Research Articles

A direct test of nitrogen and phosphorus limitation to net primary productivity in a lowland tropical wet forest

Experimental evidence for limitation of net primary productivity (NPP) by nitrogen (N) or phosphorus (P) in lowland tropical forests is rare, and the results from the few existing studies have been inconclusive. To directly test if N or P limit NPP in a lowland tropical wet forest in Costa Rica, we conducted a full factorial fertilization experiment (4 treatments x 6 replicates in 30 x 30 m plots). We focused on the influence of tree size and taxa on nutrient limitation, because in these forests a wide variety of tree functional traits related to nutrient acquisition and use are likely to regulate biogeochemical processes. After 2.7 years, a higher percentage of trees per plot increased basal area (BA) with P additions (66.45% +/- 3.28% without P vs. 76.88% +/- 3.28% with P), but there were no other community-level responses to N or P additions on BA increase, litterfall productivity, or root growth. Phosphorus additions resulted in doubled stem growth rates in small trees (5-10 cm diameter at breast height (dbh); [P < or = 0.01]) but had no effect on intermediate (10-30 cm dbh) or large trees (> 30 cm dbh). Phosphorus additions also increased the percentage of seedling survival from 59% to 78% (P < 0.01), as well as the percentage of seedlings that grew (P = 0.03), and increased leaf number (P = 0.02). Trees from Pentaclethra macroloba, the most abundant species, did not increase growth rates with fertilization (P = 0.40). In contrast, the most abundant palms (Socratea exorrhiza) had more than two times higher stem growth rates with P additions (P = 0.01). Our experiment reiterates that P availability is a significant driver of plant processes in these systems, but highlights the importance of considering different aspects of the plant community when making predictions concerning nutrient limitation. We postulate that in diverse, lowland tropical forests "heterogeneous nutrient limitation" occurs, not only driven by variability in nutrient responses among taxa, but also among size classes and potential functional groups. Heterogeneous responses to nutrient additions could lead to changes in forest structure or even diversity in the long-term, affecting rates of NPP and thus carbon cycling.

Phenotypic Responses of a Stoloniferous Clonal Plant Buchloe dactyloides to Scale-Dependent Nutrient Heterogeneity

Clonal plants could modify phenotypic responses to nutrients heterogeneously distributed both in space and time by physiological integration. It will take times to do phenotypic responses to modifications which are various in different growth periods. An optimal phenotype is reached when there is a match between nutrient conditions and foraging ability. A single plantlet of Buchloe dactyloides with two stolons was transplanted into heterogeneous nutrient conditions. One stolon grew in homogeneous nutrient patch, while the other cultured in different scales of heterogeneous nutrient patches. As compared to the other nutrient treatment, heterogeneous nutrient treatments with small scale of 25×25 cm resulted in a higher biomass, and larger number of ramets, clumps and stolons in B. dactyloides at both genet and clonal fragment levels. Significant differences of number of ramets, clumps and stolons were detected at the rapid growth stage, but not in the early stage of the experiment. Foraging ability was more efficient in heterogeneous than in homogeneous nutrient conditions as assessed by higher root mass and root to shoot ratio. Different nutrient treatments did not prompt significant differences in internode and root length. Physiological integration significantly increased biomass, but did not influence other growth or morphological characters. These results suggest that physiological integration modifies phenotypic plasticity of B. dactyloides for efficient foraging of nutrients in heterogeneous nutrient conditions. These effects are more pronounced at genet and clonal fragment levels when the patch scale is 25×25 cm. Time is a key factor when phenotypic plasticity of B. dactyloides in heterogeneous nutrient conditions is examined.

A novel morphological response of maize (Zea mays) adult roots to heterogeneous nitrate supply revealed by a split‐root experiment

Approximately 35-55% of total nitrogen (N) in maize plants is taken up by the root at the reproductive stage. Little is known about how the root of an adult plant responds to heterogeneous nutrient supply. In this study, root morphological and physiological adaptations to nitrate-rich and nitrate-poor patches and corresponding gene expression of ZmNrt2.1 and ZmNrt2.2 of maize seedlings and adult plants were characterized. Local high nitrate (LoHN) supply increased both lateral root length (LRL) and density of the treated nodal roots of adult maize plants, but only increased LRL of the treated primary roots of seedlings. LoHN also increased plant total N acquisition but not N influx rate of the treated roots, when expressed as per unit of root length. Furthermore, LoHN markedly increased specific root length (m g(-1)) of the treated roots but significantly inhibited the growth of the lateral roots outside of the nitrate-rich patches, suggesting a systemic carbon saving strategy within a whole root system. Surprisingly, local low nitrate (LoLN) supply stimulated nodal root growth of adult plants although LoLN inhibited growth of primary roots of seedlings. LoLN inhibited the N influx rate of the treated roots and did not change plant total N content. The gene expression of ZmNrt2.1 and ZmNrt2.2 of the treated roots of seedlings and adult plants was inhibited by LoHN but enhanced by LoLN. In conclusion, maize adult roots responded to nitrate-rich and nitrate-poor patches by adaptive morphological alterations and displayed carbon saving strategies in response to heterogeneous nitrate supply.

Effects of nutrient heterogeneity and competition on root architecture of spruce seedlings: implications for an essential feature of root foraging.

BackgroundWe have limited understanding of root foraging responses when plants were simultaneously exposed to nutrient heterogeneity and competition, and our goal was to determine whether and how plants integrate information about nutrients and neighbors in root foraging processes.Methodology/Principal FindingsThe experiment was conducted in split-containers, wherein half of the roots of spruce (Picea asperata) seedlings were subjected to intraspecific root competition (the vegetated half), while the other half experienced no competition (the non-vegetated half). Experimental treatments included fertilization in the vegetated half (FV), the non-vegetated half (FNV), and both compartments (F), as well as no fertilization (NF). The root architecture indicators consisted of the number of root tips over the root surface (RTRS), the length percentage of diameter-based fine root subclasses to total fine root (SRLP), and the length percentage of each root order to total fine root (ROLP). The target plants used novel root foraging behaviors under different combinations of neighboring plant and localized fertilization. In addition, the significant increase in the RTRS of 0–0.2 mm fine roots after fertilization of the vegetated half alone and its significant decrease in fertilizer was applied throughout the plant clearly showed that plant root foraging behavior was regulated by local responses coupled with systemic control mechanisms.Conclusions/SignificanceWe measured the root foraging ability for woody plants by means of root architecture indicators constructed by the roots possessing essential nutrient uptake ability (i.e., the first three root orders), and provided new evidence that plants integrate multiple forms of environmental information, such as nutrient status and neighboring competitors, in a non-additive manner during the root foraging process. The interplay between the responses of individual root modules (repetitive root units) to localized environmental signals and the systemic control of these responses may well account for the non-additive features of the root foraging process.

Growth responses of a rhizomatous herb &lt;I&gt;Bolboschoenus planiculmis&lt;/I&gt; to scale and con-trast of soil nutrient heterogeneity

Aims Spatial heterogeneity in soil nutrients is common in nature, and clonal plants are supposed to be able to better use spatially heterogeneously distributed soil nutrients. Although scale and contrast are two of the most important elements of spatial heterogeneity, few studies have tested effects of scale and contrast of soil nutrient het- erogeneity on the growth of clonal plants. Methods We conducted a greenhouse experiment in which we grew individual ramets of a rhizomatous clonal plant, Bolboschoenus planiculmis (synonym Scirpus planiculumis), in heterogeneous conditions consisting of nu- trient-rich and nutrient-poor patches. The experiment had two levels of patch scale (large vs. small patch) crossed with two levels of patch contrast (high vs. low contrast), thereby consisting of four treatments. In each treatment, the total area of the nutrient-rich patches and that of the nutrient-poor patches were the same, and the total amount of soil nutrients was also the same in all the four treatments. Important findings At both clone (whole plant) level and patch level, patch scale, contrast or their interaction did not significantly affect biomass, number of ramets, total rhizome length or number of tubers of B. planiculmis. At the patch level, however, biomass, number of ramets, total rhizome length and number of tubers of B. planiculmis were significantly greater in the nutrient-rich patches than in the nutrient-poor ones, whereas spacer length (i.e., distance between adjacent ramets) was smaller. Such effects depended on neither patch scale nor patch contrast. Therefore, when growing in environments with heterogeneous soil nutrients, B. planiculmis was able to shorten its spacer length and likely also increased branching intensity of the rhizomes, such that more ramets and tubers (i.e., potential ramets) were placed in nutrient-rich patches. These responses are generally considered one aspect of the foraging behavior of clonal plants and are thought to be adaptive. However, the patch scale or contrast designed in the present experiment could not affect the placement of the ramets in the heterogeneous environments. We presume that the responses of B. planiculmis are likely different if a wider range of scale and contrast is used and thus future studies to test effects of scale and contrast of resource heterogeneity should consider a wider range of scale and/or contrast.

Root Foraging Increases Performance of the Clonal Plant Potentilla reptans in Heterogeneous Nutrient Environments

BackgroundPlastic root-foraging responses have been widely recognized as an important strategy for plants to explore heterogeneously distributed resources. However, the benefits and costs of root foraging have received little attention.Methodology/Principal FindingsIn a greenhouse experiment, we grew pairs of connected ramets of 22 genotypes of the stoloniferous plant Potentilla reptans in paired pots, between which the contrast in nutrient availability was set as null, medium and high, but with the total nutrient amount kept the same. We calculated root-foraging intensity of each individual ramet pair as the difference in root mass between paired ramets divided by the total root mass. For each genotype, we then calculated root-foraging ability as the slope of the regression of root-foraging intensity against patch contrast. For all genotypes, root-foraging intensity increased with patch contrast and the total biomass and number of offspring ramets were lowest at high patch contrast. Among genotypes, root-foraging intensity was positively related to production of offspring ramets and biomass in the high patch-contrast treatment, which indicates an evolutionary benefit of root foraging in heterogeneous environments. However, we found no significant evidence that the ability of plastic foraging imposes costs under homogeneous conditions (i.e. when foraging is not needed).Conclusions/SignificanceOur results show that plants of P. reptans adjust their root-foraging intensity according to patch contrast. Moreover, the results show that the root foraging has an evolutionary advantage in heterogeneous environments, while costs of having the ability of plastic root foraging were absent or very small.

Exotic annuals reduce soil heterogeneity in coastal sage scrub soil chemical and biological characteristics

Exotic plant invasions alter ecosystem structure and function above- and below-ground through plant–soil feedbacks. The resistance of ecosystems to invasion can be measured by the degree of change in microbial communities and soil chemical pools and fluxes, whereas their resilience can be measured by the ability to recover following restoration. Coastal sage scrub (CSS) is one of the most highly invaded ecosystems in the US but the response of CSS soils to exotic plant invasion is little known. We examined resistance and resilience of CSS soil chemical and biological characteristics following invasion of exotic annual grasses and forbs and restoration of the native plant community. We hypothesized that invasion of exotic plant species would change biological and chemical characteristics of CSS soils by altering soil nutrient inputs. Additionally, we expected that if exotic plants were controlled and native plants were restored, native soil characteristics would recover. We sampled two locations with invaded, restored and native CSS for plant community composition, soil chemistry and microbial communities, and phospholipid fatty acid (PLFA) profiles. Communities invaded by exotic annuals were resistant to some measured parameters but not others. Extractable nitrogen pools decreased, nitrogen cycling rates increased, and microbial biomass and fungal:bacterial ratios were altered in invaded soils, and these effects were mediated by the phenological stage of the dominant plant species. The largest impact of invasion on soils was an overall reduction of spatial heterogeneity in soil nutrients, nutrient cycling and microbial communities. Restored plots tended to recover in most biotic and chemical parameters including increased resource heterogeneity compared to invaded plots, suggesting that CSS soils are resilient but not resistant to invasion.

Contribution of Root Proliferation in Nutrient-Rich Soil Patches to Nutrient Uptake and Growth of Maize

Root proliferation can be stimulated in a heterogeneous nutrient patch; however, the functions of the root proliferation in the nutrient-rich soil patches are not fully understood. In the present study, a two-year field experiment was conducted to examine the comparative effects of localized application of ammonium and phosphorus (P) at early or late stages on root growth, nutrient uptake, and biomass of maize (Zea mays L.) on a calcareous soil in an intensive farming system. Localized supply of ammonium and P had a more evident effect on shoot and root growth, and especially stimulated fine root development at the early seedling stage, with most of the maize roots being allocated to the nutrient-rich patch in the topsoil. Although localized ammonium and P supply at the late stage also enhanced the fine root growth, the plant roots in the patch accounted for a low proportion of the whole maize roots in the topsoil at the flowering stage. Compared with the early stage, fine root length in the short-lived nutrient patch decreased by 44%–62% and the shoot dry weight was not different between heterogeneous and homogeneous nutrient supply at the late growth stage. Localized supply of ammonium and P significantly increased N and P accumulation by maize at 35 and 47 days after sowing (DAS); however, no significant difference was found among the treatments at 82 DAS and the later growth stages. The increased nutrient uptake and plant growth was related to the higher proportion of root length in the localized nutrient-enriched patch. The results indicated that root proliferation in nutrient patches contributed more to maize growth and nutrient uptake at the early than late stages.

Root system size determines plant performance following short-term soil nutrient pulses

Heterogeneity in soil resources is common in natural communities, in both space and time. In contrast to substantial research into plant responses to spatial nutrient heterogeneity, little is known about the factors that influence plants responses to temporally heterogeneous nutrient supplies. We examine (1) the effects of nutrient pulse length on plant biomass allocation patterns, (2) whether a plant’s root system prior to a nutrient pulse influences the benefit a plant can gain from that pulse, and (3) how competition and nutrient pulses interact to influence plant biomass. To address these issues, we used a pot experiment with varying initial conditions (nutrient levels and competition) designed to produce a range of root system sizes in the grass Poa pratensis. We then subjected the plants to short-term (3 and 10 days long) nutrient pulses where the same total nutrients were delivered at varying rates per unit time (intensity). Plant biomass and tissue nitrogen concentration were lowest when pulses were short relative to either long pulses or continuous supply. Plants with larger root systems at the start of the nutrient pulse gained more benefit than smaller plants because final plant size relative to initial root system size decreased as pulse intensity (nutrients delivered per unit time) increased. This study has two important implications. First, since larger plants benefit disproportionately more from a nutrient pulse than smaller plants, pre-pulse conditions such as the competitive environment and resource supply are likely key determinants of a plant’s ability to capture nutrients from a pulse. Second, the importance of pre-pulse size provides a belowground mechanism to enhance the size variability among plants in a community.

Hydration dynamics promote bacterial coexistence on rough surfaces.

Identification of mechanisms that promote and maintain the immense microbial diversity found in soil is a central challenge for contemporary microbial ecology. Quantitative tools for systematic integration of complex biophysical and trophic processes at spatial scales, relevant for individual cell interactions, are essential for making progress. We report a modeling study of competing bacterial populations cohabiting soil surfaces subjected to highly dynamic hydration conditions. The model explicitly tracks growth, motion and life histories of individual bacterial cells on surfaces spanning dynamic aqueous networks that shape heterogeneous nutrient fields. The range of hydration conditions that confer physical advantages for rapidly growing species and support competitive exclusion is surprisingly narrow. The rapid fragmentation of soil aqueous phase under most natural conditions suppresses bacterial growth and cell dispersion, thereby balancing conditions experienced by competing populations with diverse physiological traits. In addition, hydration fluctuations intensify localized interactions that promote coexistence through disproportional effects within densely populated regions during dry periods. Consequently, bacterial population dynamics is affected well beyond responses predicted from equivalent and uniform hydration conditions. New insights on hydration dynamics could be considered in future designs of soil bioremediation activities, affect longevity of dry food products, and advance basic understanding of bacterial diversity dynamics and its role in global biogeochemical cycles.

Plant responses to soil heterogeneity and global environmental change.

Recent evidence suggests that soil nutrient heterogeneity, a ubiquitous feature of terrestrial ecosystems, modulates plant responses to ongoing global change (GC). However, we know little about the overall trends of such responses, the GC drivers involved, and the plant attributes affected.We synthesized literature to answer the question: Does soil heterogeneity significantly affect plant responses to main GC drivers, such as elevated atmospheric carbon dioxide concentration (CO2), nitrogen (N) enrichment and changes in rainfall regime?Overall, most studies have addressed short-term effects of N enrichment on the performance of model plant communities using experiments conducted under controlled conditions. The role of soil heterogeneity as a modulator of plant responses to elevated CO2 may depend on the plasticity in nutrient uptake patterns. Soil heterogeneity does interact with N enrichment to determine plant growth and nutrient status, but the outcome of this interaction has been found to be both synergistic and inhibitory. The very few studies published on interactive effects of soil heterogeneity and changes in rainfall regime prevented us from identifying any general pattern.We identify the long-term consequences of soil heterogeneity on plant community dynamics in the field, and the ecosystem level responses of the soil heterogeneity × GC driver interaction, as the main knowledge gaps in this area of research.In order to fill these gaps and take soil heterogeneity and GC research a step forward, we propose the following research guidelines: 1) combining morphological and physiological plant responses to soil heterogeneity with field observations of community composition and predictions from simulation models; and 2) incorporating soil heterogeneity into a trait-based response-effect framework, where plant resource-use traits are used as both response variables to this heterogeneity and GC, and predictors of ecosystem functioning.Synthesis. There is enough evidence to affirm that soil heterogeneity modulates plant responses to elevated atmospheric CO2 and N enrichment. Our synthesis indicates that we must explicitly consider soil heterogeneity to accurately predict plant responses to GC drivers.

Community-aggregated plant traits interact with soil nutrient heterogeneity to determine ecosystem functioning

Background and aims Spatial distribution of soil nutrients (soil heterogeneity) and availability have strong effects on above- and belowground plant functional traits. Although there is ample evidence on the tight links between functional traits and ecosystem functioning, the role played by soil heterogeneity and availability as modulators of such relationship is poorly known. Methods We conducted a factorial experiment in microcosmscontaining grasses,legumesand non-legumeforbs communities differing in composition to evaluate how soil heterogeneity and availability (50 and 100 mg N) affect the links between traits and ecosystem functioning. Community-aggregated specific leaf area (SLAagg) and specific root length (SRLagg )w ere measured as both relevant response traits to soil heterogeneity and availability, and significant effect traits affecting ecosystem functioning (i.e., belowground biomass, β-glucosidase and acid phosphatase activities, and in situ N availability rate). Results SRLagg was negatively and significantly associated to β-glucosidase, phosphatase and N availability rate in the high nutrient availability and heterogeneous distribution scenario. We found a significant negative relationship between SLAagg and availability rate of mineral-N under low nutrient availability conditions. Conclusions Soil heterogeneity modulated the effects of both traits and nutrient availability on ecosystem functioning. Specific root length was the key trait associated with soil nutrient cycling and belowground biomass in contrasted heterogeneous soil conditions. The inclusion of soil heterogeneity into the trait-based response-effect framework may help to scale from plant communities to the ecosystem level.

Déjections des herbivores domestiques au pâturage : caractéristiques et rôle dans le fonctionnement des prairies

Trois principaux facteurs conditionnent les caractéristiques physiques (taille des dépôts) et chimiques (teneur en eau, en nutriments…) des déjections des herbivores domestiques au pâturage : l’espèce, l’état physiologique de l’animal et la qualité de l’herbe ingérée, qui dépendra elle-même de la composition du tapis végétal et des conditions climatiques. À travers les déjections solides s’opère une restitution de matière organique et d’éléments nutritifs non assimilés vers le système sol-plante. L’importance énergétique de ce retour est telle que la dissémination des déjections est source d’hétérogénéité à l’échelle de la pâture. Cette hétérogénéité, elle même conditionnée par le comportement des animaux et les modalités de conduite, est à son tour source de diversité, tant floristique que faunistique. L’élimination de l’excrément de la surface du sol dépend des conditions climatiques et de l’activité de la faune saprocoprophage. L’activité de cette faune modifie à la fois les propriétés de la déjection et celles du sol (porosité, aération), ce qui favorise les activités microbiennes. A court terme, les déjections de grande taille ont un effet néfaste sur la végétation et entraînent un phénomène de refus de consommation de la part des bovins et des équins. A plus long terme, les déjections entraînent une augmentation de la croissance des plantes situées à proximité, effet qui peut persister jusqu’à deux ans. Si les transformations physicochimiques et les impacts des déjections sur la faune et la flore sont assez bien connus, en revanche nos connaissances sont lacunaires lorsqu’il s’agit de faire le lien entre le fonctionnement des communautés d’insectes coprophages et les fonctionnalités écosystémiques associées.

The influence of light conditions and interspecific competition on the root foraging traits and seedling growth of two tree species

Enriched nutrient patches within natural soil represent an important source of nutrients for tree growth. In the present study, pot experiments in a heterogeneous nutrient environment were conducted to investigate the influence of light conditions and interspecific competition on the root foraging traits and seedling growth of Pinus massoniana and Schima superba. The root foraging scale and the whole-seedling biomass of both species were decreased by shading. The result of this treatment was a lower sensitivity to nutrient heterogeneity in plants that underwent the shading treatment than in plants that were exposed to full-light conditions. The above-ground biomass and whole-seedling biomass of S. superba were not affected by competition with P. massoniana. In contrast, the above-ground biomass and whole-seedling biomass of P. massoniana were negatively affected by competition with S. superba. The more rapid rate of root extension and the more efficient resource uptake of S.superba appear to explain this effect. The species-specific patterns of the influence of environmental factors on foraging ability and seedling growth should be given thorough consideration and should be applied to afforestation and to the management of tree plantations.

格氏栲天然林土壤养分空间异质性

The spatial heterogeneity of soil nutrients and soil pH in Castanopsis kawakamii natural forest of Sanming City,Fujian Province was studied by the method of Geostatistics coupled with Geographical Information System(GIS) technique.The experimental results showed that pH had a weak variation,and the soil nutrients exhibited medium spatial variability.The variation intensity of soil nutrients and soil pH ranked from high to low as follows: available phosphorous(43.54%),available potassium(32.84%),hydrolyzable nitrogen(31.03%),total nitrogen(29.41%),total potassium(25.25%),total phosphorous(22.14%),pH(3.39%).In terms of different soil layers,the value of pH and total potassium raised with increase in soil depth,while total nitrogen,total phosphorous,hydrolyzable nitrogen,available phosphorous,available potassium reduced with increasing soil depth.Analysis of variance showed that soil nutrients and soil pH reached an extremely significant levels.Parameters of theoretical variogram models for soil nutrients and soil pH showed that pH,total potassium and available phosphorous fitted exponential model,total nitrogen fitted Gaussian model,total phosphorous and hydrolyzable nitrogen can be best described by spherical model and available potassium was well fitted by linear model.The effective ranges of total potassium and available potassium were 1806 m and 549 m,respectively,while the values for pH,total nitrogen,total phosphorous,hydrolyzable nitrogen and available phosphorous were 267 m,130 m,120 m,182 m,and 117 m,respectively.In terms of spatial structure,pH,total nitrogen,total phosphorous,hydrolyzable nitrogen and available phosphorous showed high spatial autocorrelation,which was greatly affected by soil structural factors,such as stand structure,topography,microhabitat of forest gap and soil aggregate structure.Total potassium indicated moderate spatial autocorrelation,which was resulted from both soil structural factors and random factors.Available potassium indicated weak spatial autocorrelation.The spatial heterogeneity of available potassium was primarily affected by random factors,such as site preparation,change of land use,the rapid development of the ecotourism,the planning and construction of scenic areas.The spatial distributions of soil nutrients had different characteristics.Total nitrogen and total phosphorous were increased from south to north.The total potassium emerged circle distribution with low in the central area and high in the marginal area.Soil pH showed a decreasing trend from southwest to northeast,presenting a band distribution.Hydrolyzable nitrogen decreased from northeast to southwest.Available phosphorous increased gradually from southeast to northwest.From southwest to northeast,available potassium content increased firstly and then decreased with two high value areas at the corner of northwest and southeast.The results may provide reference for improving the sampling design and for spatial interpolation and map of soil nutrients at a regional scale.It also may give a scientific foundation for sustainable use of soil and the recovery and reconstruction of C.kawakamii natural forest.

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

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. 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. 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. 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.

Microscale patchiness leads to large and important intraspecific internal nutrient heterogeneity in phytoplankton

Phytoplankton stoichiometry or nutrient content has been shown to vary in a number of dimensions (species, condition, time, space), but the heterogeneity within a species at a given time and location, and the underlying mechanisms and importance have not been explored. There are a number of mechanisms that can create intraspecific heterogeneity, and theory suggests it can affect the population growth rate. We studied heterogeneity in P content of the freshwater diatom Cyclotella meneghiniana in the Charles River in Boston. Single-cell observations using synchrotron-based X-ray fluorescence show that the nutrient status varies from P-starved to P-replete. We simulate individual cells using an agent-based model that accounts for a number of mechanisms that can create heterogeneity, including surface area–based uptake, mortality differentiation, stochastic biological variability in states and behavior, macroscale mixing, and microscale nutrient patch encounter. By performing a number of simulations with various mechanisms turned on/off and comparing to data, we conclude that the heterogeneity is mostly due to microscale patchiness (85%). We explore the importance of accounting for heterogeneity in models by performing a simulation with the growth rate based on the population-average internal nutrient, as is done in conventional population-level models. This shows that ignoring heterogeneity increases the population growth rate by a factor of 1.47. To account for different heterogeneity in the laboratory and field, population-level ecosystem models should reduce maximum growth rates. The magnitude of this correction depends on local conditions, and in our case, it is a factor of 0.72.

Soil nutrient patchiness and plant genotypes interact on the production potential and decomposition of root and shoot litter: evidence from short-term laboratory experiments with Triticum aestivum

Aims The purpose of this study was to test the hypotheses that soil nutrient patchiness can differentially benefit the decomposition of root and shoot litters and that this facilitation depends on plant genotypes. Methods We grew 15 cultivars (i.e. genotypes) of winter wheat (Triticum aestivum L.) under uniform and patchy soil nutrients, and contrasted their biomass and the subsequent mass, carbon (C) and nitrogen (N) dynamics of their root and shoot litters. Results Under equal amounts of nutrients, patchy distribution increased root biomass and had no effects on shoot biomass and C:N ratios of roots and shoots. Roots and shoots decomposed more rapidly in patchy nutrients than in uniform nutrients, and reductions in root and shoot C:N ratios with decomposition were greater in patchy nutrients than uniform nutrients. Soil nutrient patchiness facilitated shoot decomposition more than root decomposition. The changes in C:N ratios with decomposition were correlated with initial C:N ratios of litter, regardless of roots or shoots. Litter potential yield, quality and decomposition were also affected by T. aestivum cultivars and their interactions with nutrient patchiness. Conclusions Soil nutrient patchiness can enhance C and N cycling and this effect depends strongly on genotypes of T. aestivum. Soil nutrient heterogeneity in plant communities also can enhance diversity in litter decomposition and associated biochemical and biological dynamics in the soil.

Interactive effects of nutrient heterogeneity and competition: implications for root foraging theory?

Summary1. Plants are known to respond to heterogeneous distribution of nutrients in the soil, and they also respond to the presence of neighbouring roots. However, it is unclear whether plants are able to distinguish between these factors and adjust their root responses accordingly.2. We investigated whether the simultaneous response to nutrient heterogeneity and competition could be predicted from the responses to these factors separately. As a null model, we hypothesized that the responses to nutrients and competition are additive and thus no interactions occur. We performed a short‐term competition experiment in the greenhouse with two floodplain species in homogeneous and heterogeneous conditions. The consequences of different root distributions for nutrient uptake were tested using 15N pulse‐labelling.3. Both species responded to nutrient heterogeneity by investing significantly more roots in the nutrient‐rich patch, and both species showed a significant reduction in root growth in response to competition, albeit that the reduction was much more pronounced for the grass species. For Rumex palustris, the effects of heterogeneity and competition were additive. However, the response to nutrient heterogeneity of Agrostis stolonifera was reversed by competition: instead of proliferating in the nutrient‐rich patch, it significantly increased root investments in the ‘empty’ (nutrient‐poor) patches. As the partitioning of total N was less asymmetric than 15N uptake from the nutrient‐rich patch, it appears that these altered root investments of A. stolonifera in the ‘empty’ patches have also been functional with respect to compensating N uptake.4. Our results suggest that root responses to nutrient distribution in a competitive environment depend on the competitive strength of the neighbouring species. The foraging response of the superior species (R. palustris) was hardly affected, but that of the inferior species (A. stolonifera) was greatly inhibited and even reversed by competition: instead of proliferating in the nutrient‐rich patch, it increased root growth and foraging activity in less favourable patches. Incorporating competitive hierarchy into root foraging studies may help to explain the ambiguous results found in previous studies.

Optimal root proliferation strategies: the roles of nutrient heterogeneity, competition and mycorrhizal networks

Background and Aims Plants proliferate roots in order to acquire nutrients, typically contending with heterogeneous resources and competing neighbours. A mathematical model was developed to identify optimal root proliferation strategies in patchy nutrient environments. The impact of joining mycorrhizal networks was also assessed. Methods A simple model of growth and competition in one spatial dimension was implemented within a genetic algorithm to obtain optimal proliferation strategies under different scenarios of resource distribution, and in the presence or absence of local competition and large-scale mycorrhizal networks. Results A strong proliferation response emerged for isolated plants in heterogeneous environments with low resources, and also for plants growing in competition. Even in statistically homogeneous environments, the presence of competition conferred a selective advantage to plants proliferating in the direction of the most recently acquired patch. In the presence of mycorrhizal networks, the optimal strategy switched from symbiosis to proliferation driven growth as the relative cost of acquiring resources through the networks increased. Conclusions The optimal proliferation response in a given scenario was governed by a hierarchy of factors: resource levels and distribution; the presence or absence of competition; and the marginal benefit of obtaining resources via symbiotic relationships with mycorrhizas.