The role of above-ground competition and nitrogen vs. phosphorus enrichment in seedling survival of common European plant species of semi-natural grasslands.
ABSTRACT: Anthropogenic activities have severely altered fluxes of nitrogen and phosphorus in ecosystems worldwide. In grasslands, subsequent negative effects are commonly attributed to competitive exclusion of plant species following increased above-ground biomass production. However, some studies have shown that this does not fully account for nutrient enrichment effects, questioning whether lowering competition by reducing grassland productivity through mowing or herbivory can mitigate the environmental impact of nutrient pollution. Furthermore, few studies so far discriminate between nitrogen and phosphorus pollution. We performed a full factorial experiment in greenhouse mesocosms combining nitrogen and phosphorus addition with two clipping regimes designed to relax above-ground competition. Next, we studied the survival and growth of seedlings of eight common European grassland species and found that five out of eight species showed higher survival under the clipping regime with the lowest above-ground competition. Phosphorus addition negatively affected seven plant species and nitrogen addition negatively affected four plant species. Importantly, the negative effects of nutrient addition and higher above-ground competition were independent of each other for all but one species. Our results suggest that at any given level of soil nutrients, relaxation of above-ground competition allows for higher seedling survival in grasslands. At the same time, even at low levels of above-ground competition, nutrient enrichment negatively affects survival as compared to nutrient-poor conditions. Therefore, although maintaining low above-ground competition appears essential for species' recruitment, for instance through mowing or herbivory, these management efforts are likely to be insufficient and we conclude that environmental policies aimed to reduce both excess nitrogen and particularly phosphorus inputs are also necessary.
Project description:Vachellia sieberiana fixes atmospheric nitrogen (N) and distributes it back into ecosystems. We hypothesize that biological nitrogen fixation in this plant species is limited by competition from the invasive shrub, Chromolaena odorata. Competition would therefore result in the legume plant switching its limited nitrogen (N) sources in phosphorus-poor soils in savannah ecosystems when resources have to be shared. This study investigated the different patterns of N use and growth costs by a native and an introduced leguminous shrubby species. We propose that the two species sharing the same environment might result in competition. The competitive effect would induce in the indigenous legume to better utilize atmospheric-derived N modifying plant growth kinetics and plant mineral concentrations. Seedlings of V. sieberiana were cultivated in natural soil inoculum with low levels of phosphorus (mg L-1 ± SE) of 3.67 ± 0.88. The experiments were divided into two treatments where (i) seedlings of V. sieberiana were subjected to competition by cultivating them together with seedlings of C. odorata, and (ii) seedlings of V. sieberiana were cultivated independently. Although V. sieberiana was subjected to competition, the N2-fixing bacteria that occupied the nodules was Mesorhizobium species, similar to plants not subjected to competition. Total plant biomass was similar between treatments although V. sieberiana plants subjected to competition accumulated more below-ground biomass and showed higher carbon construction costs than plants growing individually. Total plant phosphorus and nitrogen decreased in seedlings of V. sieberiana under competition, whereas no differences were observed in percent N derived from the atmosphere (%NDFA) between treatments. The specific nitrogen utilization rate (SNUR) was higher in V. sieberiana plants subjected to competition while specific nitrogen absorption rate (SNAR) showed the opposite response. Vachellia sieberiana is highly adapted to nutrient-poor savannah ecosystems and can withstand competition from invasive shrubs by utilizing both atmospheric and soil nitrogen sources.
Project description:Insect herbivores can shift the composition of a plant community, but the mechanism underlying such shifts remains largely unexplored. A possibility is that insects alter the competitive symmetry between plant species. The effect of herbivory on competition likely depends on whether the plants are subjected to aboveground or belowground herbivory or both, and also depends on soil nitrogen levels. It is unclear how these biotic and abiotic factors interactively affect competition. In a greenhouse experiment, we measured competition between two coexisting grass species that respond differently to nitrogen deposition: Dactylis glomerata L., which is competitively favoured by nitrogen addition, and Festuca rubra L., which is competitively favoured on nitrogen-poor soils. We predicted: (1) that aboveground herbivory would reduce competitive asymmetry at high soil nitrogen by reducing the competitive advantage of D. glomerata; and (2), that belowground herbivory would relax competition at low soil nitrogen, by reducing the competitive advantage of F. rubra. Aboveground herbivory caused a 46% decrease in the competitive ability of F. rubra, and a 23% increase in that of D. glomerata, thus increasing competitive asymmetry, independently of soil nitrogen level. Belowground herbivory did not affect competitive symmetry, but the combined influence of above- and belowground herbivory was weaker than predicted from their individual effects. Belowground herbivory thus mitigated the increased competitive asymmetry caused by aboveground herbivory. D. glomerata remained competitively dominant after the cessation of aboveground herbivory, showing that the influence of herbivory continued beyond the feeding period. We showed that insect herbivory can strongly influence plant competitive interactions. In our experimental plant community, aboveground insect herbivory increased the risk of competitive exclusion of F. rubra. Belowground herbivory appeared to mitigate the influence of aboveground herbivory, and this mechanism may play a role for plant species coexistence.
Project description:While mowing-induced changes in plant traits and their effects on ecosystem functioning in semi-arid grassland are well studied, the relations between plant size and nutrient strategies are largely unknown. Mowing may drive the shifts of plant nutrient limitation and allocation. Here, we evaluated the changes in nutrient stoichiometry and allocation with variations in sizes of Leymus chinensis, the dominant plant species in Inner Mongolia grassland, to various mowing frequencies in a 17-yr controlled experiment. Affected by mowing, the concentrations of nitrogen (N), phosphorus (P), and carbon (C) in leaves and stems were significantly increased, negatively correlating with plant sizes. Moreover, we found significant trade-offs between the concentrations and accumulation of N, P, and C in plant tissues. The N:P ratios of L. chinensis aboveground biomass, linearly correlating with plant size, significantly decreased with increased mowing frequencies. The ratios of C:N and C:P of L. chinensis individuals were positively correlated with plant size, showing an exponential pattern. With increased mowing frequencies, L. chinensis size was correlated with the allocation ratios of leaves to stems of N, P, and C by the tendencies of negative parabola, positive, and negative linear. The results of structure equation modeling showed that the N, P, and C allocations were co-regulated by biomass allocation and nutrient concentration ratios of leaves to stems. In summary, we found a significant decoupling effect between plant traits and nutrient strategies along mowing frequencies. Our results reveal a mechanism for how long-term mowing-induced changes in concentrations, accumulations, ecological stoichiometry, and allocations of key elements are mediated by the variations in plant sizes of perennial rhizome grass.
Project description:<h4>Background and aims</h4>During their domestication, maize, bean and squash evolved in polycultures grown by small-scale farmers in the Americas. Polycultures often overyield on low-fertility soils, which are a primary production constraint in low-input agriculture. We hypothesized that root architectural differences among these crops causes niche complementarity and thereby greater nutrient acquisition than corresponding monocultures.<h4>Methods</h4>A functional-structural plant model, SimRoot, was used to simulate the first 40 d of growth of these crops in monoculture and polyculture and to determine the effects of root competition on nutrient uptake and biomass production of each plant on low-nitrogen, -phosphorus and -potassium soils.<h4>Key results</h4>Squash, the earliest domesticated crop, was most sensitive to low soil fertility, while bean, the most recently domesticated crop, was least sensitive to low soil fertility. Nitrate uptake and biomass production were up to 7 % greater in the polycultures than in the monocultures, but only when root architecture was taken into account. Enhanced nitrogen capture in polycultures was independent of nitrogen fixation by bean. Root competition had negligible effects on phosphorus or potassium uptake or biomass production.<h4>Conclusions</h4>We conclude that spatial niche differentiation caused by differences in root architecture allows polycultures to overyield when plants are competing for mobile soil resources. However, direct competition for immobile resources might be negligible in agricultural systems. Interspecies root spacing may also be too large to allow maize to benefit from root exudates of bean or squash. Above-ground competition for light, however, may have strong feedbacks on root foraging for immobile nutrients, which may increase cereal growth more than it will decrease the growth of the other crops. We note that the order of domestication of crops correlates with increasing nutrient efficiency, rather than production potential.
Project description:Background:Oviposition decisions are critical to the fitness of herbivorous insects and are often impacted by the availability and condition of host plants. Monarch butterflies (Danaus plexippus) rely on milkweeds (Asclepias spp.) for egg-laying and as food for larvae. Previous work has shown that monarchs prefer to oviposit on recently regrown plant tissues (after removal of above-ground biomass) while larvae grow poorly on plants previously damaged by insects. We hypothesized that these effects may depend on the life-history strategy of plants, as clonal and non-clonal milkweed species differ in resource allocation and defense strategies. Methodology/Principal Findings:We first confirmed butterfly preference for regrown tissue in a field survey of paired mowed and unmowed plots of the common milkweed A. syriaca. We then experimentally studied the effects of plant damage (comparing undamaged controls to plants clipped and regrown, or damaged by insects) on oviposition choice, larval performance, and leaf quality of two closely related clonal and non-clonal species pairs: (1) A. syriaca and A. tuberosa, and (2) A. verticillata and A. incarnata. Clonal and non-clonal species displayed different responses to plant damage, impacting the proportions of eggs laid on plants. Clonal species had similar mean proportions of eggs on regrown and control plants (?35-40% each), but fewer on insect-damaged plants (?20%). Meanwhile non-clonal species had similar oviposition on insect-damaged and control plants (20-30% each) but more eggs on regrown plants (40-60%). Trait analyses showed reduced defenses in regrown plants and we found some evidence, although variable, for negative effects of insect damage on subsequent larval performance. Conclusions/Significance:Overall, non-clonal species are more susceptible and preferred by monarch butterflies following clipping, while clonal species show tolerance to clipping and induced defense to insect herbivory. These results have implications for monarch conservation strategies that involve milkweed habitat management by mowing. More generally, plant life-history may mediate growth and defense strategies, explaining species-level variation in responses to different types of damage.
Project description:Predicting species responses to climate change and land use practices requires understanding both the direct effects of environmental factors as well as the indirect effects mediated by changes in belowground and aboveground competition. Belowground root competition from surrounding vegetation and aboveground light competition are two important factors affecting seedling establishment. However, few studies have jointly examined the effect of belowground root and light competition on seedling establishment, especially under long-term nitrogen addition and mowing. Here, we examined how belowground root competition from surrounding vegetation and aboveground light competition affect seedling establishment within a long-term nitrogen addition and mowing experiment. Seedlings of two grasses (Stipa krylovii and Cleistogenes squarrosa) were grown with and without belowground root competition under control, nitrogen addition, and mowing treatments, and their growth characteristics were monitored. The seedlings of the two grasses achieved higher total biomass, height, mean shoot and root mass, but a lower root/shoot ratio in the absence than in the presence of belowground root competition. Nitrogen addition significantly decreased shoot biomass, root biomass, and the survival of the two grasses. Regression analyses revealed that the biomass of the two grass was strongly negatively correlated with net primary productivity under belowground root competition, but with the intercept photosynthetic active radiation in the absence of belowground root competition. This experiment demonstrates that belowground root competition can alter the grass seedling establishment response to light in a long-term nitrogen addition and mowing experiment. Graphical Abstract Effects of light competition and belowground competition on the seedling of S. krylovii and C. squarrosa under the control, mowing, and N addition treatment.
Project description:Nutrient enrichment, particularly nitrogen, is an important determinant of plant community productivity, diversity and invasibility in a wetland ecosystem. It may contribute to increasing colonization and dominance of invasive species, such as Phragmites australis, especially during wetland restoration. Providing native species a competitive advantage over invasive species, manipulating soil nutrients (nitrogen) may be an effective strategy to control the invasive species and that management tool is essential to restore the degraded ecosystems. Therefore, we examined competition between Phragmites australis and Melaleuca ericifolia in a greenhouse setting with activated carbon (AC) treatments, followed by cutting of Phragmites shoots in nutrient-rich soils. Additionally, we evaluated the effect of AC on plant-free microcosms in the laboratory, to differentiate direct effects of AC on soil microbial functions from indirect effects. Overall, the objective was to test whether lowering nitrogen might be an effective approach for reducing Phragmites invasion in the wetland. The AC reduced Phragmites total biomass more significantly in repeated cut regime (57%) of Phragmites shoots compared to uncut regime (39%). Conversely, it increased Melaleuca total biomass by 41% and 68% in uncut and repeated cut regimes, respectively. Additionally, AC decreased more total nitrogen in above-ground biomass (41 to 55%) and non-structural carbohydrate in rhizome (21 to 65%) of Phragmites, and less total nitrogen reduction in above-ground biomass (25 to 24%) of Melaleuca in repeated cut compared to uncut regime. The significant negative correlation between Phragmites and Melaleuca total biomass was observed, and noticed that Phragmites acquired less biomass comparatively than Melaleuca in AC-untreated versus AC-treated pots across the cutting frequency. AC also caused significant changes to microbial community functions across Phragmites populations, namely nitrogen mineralization, nitrification, nitrogen microbial biomass and dehydrogenase activity (P???0.05) that may potentially explain changes in plant growth competition between Phragmites and Melaleuca. The overall effects on plant growth, however, may be partially microbially mediated, which was demonstrated through soil microbial functions. Results support the idea that reducing community vulnerability to invasion through nutrient (nitrogen) manipulations by AC with reducing biomass of invasive species may provide an effective strategy for invasive species management and ecosystem restoration.
Project description:<h4>Background and aims</h4>Knowledge of plant resource acquisition strategies is crucial for understanding the mechanisms mediating the responses of ecosystems to external nitrogen (N) input. However, few studies have considered the joint effects of above-ground (light) and below-ground (nutrient) resource acquisition strategies in regulating plant species responses to N enrichment. Here, we quantified the effects of light and non-N nutrient acquisition capacities on species relative abundance in the case of extra N input.<h4>Methods</h4>Based on an N-manipulation experiment in a Tibetan alpine steppe, we determined the responses of species relative abundances and light and nutrient acquisition capacities to N enrichment for two species with different resource acquisition strategies (the taller Stipa purpurea, which is colonized by arbuscular mycorrhizal fungi, and the shorter Carex stenophylloides, which has cluster roots). Structural equation models were developed to explore the relative effects of light and nutrient acquisition on species relative abundance along the N addition gradient.<h4>Key results</h4>We found that the relative abundance of taller S. purpurea increased with the improved light acquisition along the N addition gradient. In contrast, the shorter C. stenophylloides, with cluster roots, excelled in acquiring phosphorus (P) so as to elevate its leaf P concentration under N enrichment by producing large amounts of carboxylate exudates that mobilized moderately labile and recalcitrant soil P forms. The increased leaf P concentration of C. stenophylloides enhanced its light use efficiency and promoted its relative abundance even in the shade of taller competitors.<h4>Conclusions</h4>Our findings highlight that the combined effects of above-ground (light) and below-ground (nutrient) resources rather than light alone (the prevailing perspective) determine the responses of grassland community structure to N enrichment.
Project description:Submerged macrophytes play an important role in maintaining good water quality in shallow lakes. Yet extensive stands easily interfere with various services provided by these lakes, and harvesting is increasingly applied as a management measure. Because shallow lakes may possess alternative stable states over a wide range of environmental conditions, designing a successful mowing strategy is challenging, given the important role of macrophytes in stabilizing the clear water state. In this study, the integrated ecosystem model PCLake is used to explore the consequences of mowing, in terms of reducing nuisance and ecosystem stability, for a wide range of external nutrient loadings, mowing intensities and timings. Elodea is used as a model species. Additionally, we use PCLake to estimate how much phosphorus is removed with the harvested biomass, and evaluate the long-term effect of harvesting. Our model indicates that mowing can temporarily reduce nuisance caused by submerged plants in the first weeks after cutting, particularly when external nutrient loading is fairly low. The risk of instigating a regime shift can be tempered by mowing halfway the growing season when the resilience of the system is highest, as our model showed. Up to half of the phosphorus entering the system can potentially be removed along with the harvested biomass. As a result, prolonged mowing can prevent an oligo-to mesotrophic lake from becoming eutrophic to a certain extent, as our model shows that the critical nutrient loading, where the lake shifts to the turbid phytoplankton-dominated state, can be slightly increased.
Project description:Biomass allocation affects the ability of plants to acquire resources and nutrients; a limited allocation of nutrients, such as nitrogen and phosphorus, affects ecological processes. However, little research has been conducted on how plant allocation patterns change and on the trade-offs involved in allocation strategies when microhabitat gradients exist. We selected a 3.6 km transect in the Ebinur Lake Wetland Natural Reserve of Xinjiang, China, to investigate the relationships between plant traits (biomass and N and P concentrations) of herbaceous plants and environmental factors (soil moisture, salinity and nutrient content), and to determine the allometric scaling of biomass and stoichiometric traits between the above- and below-ground plant parts. The results show that the biomass and stoichiometric traits of plants reflected both the change of micro-environment and the natural characteristics of plants. With a decrease of the soil water availability and salinity, above- and below-ground N and P concentrations decrease gradually; scaling relationships exist between above- and below-ground plant parts, for biomass and N and P concentrations. Biomass allocation is influenced by soil nutrient ratios, and the allocation strategy tended to be conserved for N and variable for P. Second, the scaling relationships also show interspecific differences; all scaling exponents of <i>Suaeda prostrata</i> are larger than for other species and indicate a 'tolerance' strategy, while other species tend to increase the below-ground biomass and N and P concentrations, i.e. a 'capture' strategy.