Correction to: Comparing the impacts of an invasive grass on nitrogen cycling and ammonia-oxidizing prokaryotes in high-nitrogen forests,open fields,and wetlands

Plant and Soil - Crop genotypes may respond differently to various physical soil conditions. The objective of this study was to investigate the responses of the root architectures of two maize...     

Positive feedbacks between decomposition and soil nitrogen availability along fertility gradients

Background and aims

We determined the relationship between site N supply and decomposition rates with respect to controls exerted by environment, litter chemistry, and fungal colonization.

Methods

Two reciprocal transplant decomposition experiments were established, one in each of two long-term experiments in oak woodlands in Minnesota, USA: a fire frequency/vegetation gradient, along which soil N availability varies markedly, and a long-term N fertilization experiment. Both experiments used native Quercus ellipsoidalis E.J. Hill and Andropogon gerardii Vitman leaf litter and either root litter or wooden dowels.

Results

Leaf litter decay rates generally increased with soil N availability in both experiments while belowground litter decayed more slowly with increasing soil N. Litter chemistry differed among litter types, and these differences had significant effects on belowground (but not aboveground) decay rates and on aboveground litter N dynamics during decomposition. Fungal colonization of detritus was positively correlated with soil fertility and decay rates.

Conclusions

Higher soil fertility associated with low fire frequency was associated with greater leaf litter production, higher rates of fungal colonization of detritus, more rapid leaf litter decomposition rates, and greater N release in the root litter, all of which likely enhance soil fertility. During decomposition, both greater mass loss and litter N release provide mechanisms through which the plant and decomposer communities provide positive feedbacks to soil fertility as ultimately driven by decreasing fire frequency in N-limited soils and vice versa.     

Soil nitrogen cycling rates in low arctic shrub tundra are enhanced by litter feedbacks

Shrub growth has increased across the Arctic in recent decades and is strongly limited by soil nitrogen (N) availability. In order to understand the role of N in controlling shrub growth, we compared N-cycling in tall birch (Betula glandulosa) and surrounding dwarf birch hummock vegetation on similar soils in a Canadian low arctic site. Stable isotope tracer analysis revealed N pools and cycling rates were ～3 times larger and faster in the tall birch ecosystem in the late growing season, just prior to leaf senescence. Gross NH ₄ ⁺ -N production rates in these ecosystems correlated positively with larger pools and production rates of dissolved soil C and N, higher quality litter inputs and lower soil C. Analyses of the soil microbial community in both ecosystems indicated similar fungal dominance (epifluorescence microscopy) and similar compositions of the principal fungal or bacterial phylotypes (denaturing gradient gel electrophoresis). Together, these results strongly suggest that vegetation feedbacks associated with larger inputs of higher quality litter promote rapid soil N-cycling and enhanced shrub growth in tall birch tundra. We conclude that these litter-related feedbacks during summer may be as important as snow-shrub feedbacks in maintaining and promoting differences in shrub growth across the arctic landscape.     

The cycling of organic nitrogen through the atmosphere

Atmospheric organic nitrogen (ON) appears to be a ubiquitous but poorly understood component of the atmospheric nitrogen deposition flux. Here, we focus on the ON components that dominate deposition and do not consider reactive atmospheric gases containing ON such as peroxyacyl nitrates that are important in atmospheric nitrogen transport, but are probably not particularly important in deposition. We first review the approaches to the analysis and characterization of atmospheric ON. We then briefly summarize the available data on the concentrations of ON in both aerosols and rainwater from around the world, and the limited information available on its chemical characterization. This evidence clearly shows that atmospheric aerosol and rainwater ON is a complex mixture of material from multiple sources. This synthesis of available information is then used to try and identify some of the important sources of this material, in particular, if it is of predominantly natural or anthropogenic origin. Finally, we suggest that the flux of ON is about 25 per cent of the total nitrogen deposition flux.     

Novel plant-soil feedbacks drive adaption of invasive plants to soil legacies of native plants under nitrogen deposition

Plant and Soil - Soil legacies mediate interactions between native and introduced plants, contributing to both invasion and biotic resistance to invasion. Given that nitrogen deposition can promote...     

Microbial-mediated feedbacks of leaf litter on invasive plant growth and interspecific competition

Background and Aims

In line with the Stress Gradient Hypothesis, studies of facilitation have tended to focus on plant–plant interactions (biotic nurses), while the relative role of abiotic nurses has been little studied. We assessed the role of biotic and abiotic nurses, and their interaction, on soil enhancement and the consequential performance of a native annual grass, Dactyloctenium radulans.

Methods

We used a growth chamber study with two levels of water application to compare the performance of D. radulans growing in soil from foraging pits of the Short-beaked echidna (Tachyglossus aculeatus; abiotic nurse) and non-pit soil from either under tree canopies (biotic nurse) or surrounding open areas.

Results

All measures of plant performance were more pronounced under the high than the low water treatment. The greatest differences between pit and surface Microsites occurred under the low water application, reinforcing our view that facilitatory effects are greater in resource-limited environments. Despite tree canopy soil having greater N, there was no significant effect on plant performance, nor any significant interaction with Microsite.

Conclusions

Our study provides strong evidence that foraging pits enhance soil properties and this soil, in turn, facilitates plant growth; and supports previous work documenting the positive effect of nurse-protégé interactions under greater levels of abiotic stress.     

Contrasting impacts of grass species on nitrogen cycling in a grazed Sudanian savanna

We investigated the impact of perennial and annuals grass species on nitrogen cycling in a Sudanian savanna of Burkina Faso. We also analysed how the local context in terms of grazing and soil properties modifies these impacts. We selected four plots differing both by the intensity of grazing by cattle and soil depth, and used soil and grass biomass ¹⁵N as integrative indicators of N cycle. If perennials are able to foster a more efficient nitrogen cycling there should be lower ¹⁵N abundances in their biomass and soil. If soil depth and cattle pressure significantly modify nitrogen fluxes, soil depth and cattle pressure should influence ¹⁵N signatures. Our results suggest that perennial grasses are more conservative for nitrogen (inhibition of nitrification, less leaching via a perennial root system, slower cycling). The increase in leaf δ¹⁵N with N concentration is steeper in Loudetia togoensis than in the three other grasses. No significant difference was found between the ¹⁵N signatures of the four plots. Our results on ¹⁵N signatures and the fact that perennial grasses are much more abundant in the plots that are less grazed and have deeper soils, confirm that the switch from perennial to annual grasses is linked to a degradation in soil fertility and pasture quality. This suggests that ¹⁵N signatures can be used as indicators of fertility.     

Small population size limits reproduction in an invasive grass through both demography and genetics

Small populations of founding individuals or survivors of incomplete management programs often represent critical transitions in biological invasions. Theory predicts that population size affects reproduction and, consequently, a population’s expansion, but there are few empirical tests, and fewer that account for the reduced genetic diversity that often accompanies small population size. We created experimental small populations of invasive ryegrass (Lolium multiflorum) with population size varying independently from genetic diversity. Treatment independence was achieved by cloning plants to increase population size without changing diversity. Plant fitness was measured as the proportion of florets producing a seed. We analyzed the effects of population size, genetic diversity, and their interaction using ANCOVAs, one of which accounted for variation in individual plant growth. As predicted, smaller populations produced significantly lower proportion seed set. Low genetic diversity also reduced seed set, but this was best interpreted as part of a significant interaction with population size. Specifically, the effect of population size on the proportion seed set was over five times larger for populations in the medium genetic diversity treatment than the highest diversity treatment, and 6.7 times larger for populations with the lowest level of diversity. Population size variation had biologically meaningful consequences, as the rate of seed set within the low diversity treatment increased by 80 % with increasing population size. The results indicate that both the demographics and genetics of populations can influence reproduction and invasive potential, and must be considered when assessing risk and designing management plans for invasive plants.     

Nutrient feedbacks to soil heterotrophic nitrogen fixation in forests

Multiple nutrient cycles regulate biological nitrogen (N) fixation in forests, yet long-term feedbacks between N-fixation and coupled element cycles remain largely unexplored. We examined soil nutrients and heterotrophic N-fixation across a gradient of 24 temperate conifer forests shaped by legacies of symbiotic N-fixing trees. We observed positive relationships among mineral soil pools of N, carbon (C), organic molybdenum (Mo), and organic phosphorus (P) across sites, evidence that legacies of symbiotic N-fixing trees can increase the abundance of multiple elements important to heterotrophic N-fixation. Soil N accumulation lowered rates of heterotrophic N-fixation in organic horizons due to both N inhibition of nitrogenase enzymes and declines in soil organic matter quality. Experimental fertilization of organic horizon soil revealed widespread Mo limitation of heterotrophic N-fixation, especially at sites where soil Mo was scarce relative to C. Fertilization also revealed widespread absence of P limitation, consistent with high soil P:Mo ratios. Responses of heterotrophic N-fixation to added Mo (positive) and N (negative) were correlated across sites, evidence that multiple nutrient controls of heterotrophic N-fixation were more common than single-nutrient effects. We propose a conceptual model where symbiotic N-fixation promotes coupled N, C, P, and Mo accumulation in soil, leading to positive feedback that relaxes nutrient limitation of overall N-fixation, though heterotrophic N-fixation is primarily suppressed by strong negative feedback from long-term soil N accumulation.     

Non-native competitive perennial grass impedes the spread of an invasive annual grass

Invasive plants are degrading wildlands around the globe by displacing native species, reducing biodiversity, and altering ecological functions. The current approach of applying herbicides to invasive plants in wildlands has not been effective at curtailing their expansion and, in certain circumstances, may do more harm than good. Preventing the spread of invasive species has been identified as an important strategy to protect wildlands. However, few prevention strategies have actually been tested. We hypothesized that establishing competitive vegetation next to infestations would increase the biotic resistance of the plant community to invasion and decrease the invasive species propagule pressure beyond the competitive vegetation. To evaluate this, we established twelve competitive vegetation barriers in front of invasive annual grass, Taeniatherum caput-medusae (L.) Nevski, infestations. The non-native perennial grass Agropyron desertorum (Fisch. ex Link) Schult. was seeded into plant communities adjacent to the infestations to create the competitive vegetation barriers. Soil nutrient concentrations and the spread of T. caput-medusae were compared between where A. desertorum was seeded and not seeded (control treatment) 3 years after treatment. Less T. caput-medusae and lower soil ammonium and potassium concentrations in the competitive vegetation barrier than control treatment (P ≤ 0.01) suggest that establishing competitive vegetation increased the biotic resistance of the plant communities to invasion. Taeniatherum caput-medusae cover and density in the plant communities protected by the competitive vegetation barrier (locales across the barriers from the infestations) were ~42- and 47-fold less, respectively, than unprotected plant communities (P < 0.01). This suggests that invasive plant propagule pressure was decreased in the plant communities protected by competitive vegetation barriers. The establishment of competitive vegetation around infestations may be an effective strategy to prevent or at least reduce the spread of invasive plant species.     

Constricted growth of grass roots through rigid pores

Summary An examination of the mechanical interaction between elongating roots and rigid pores of precisely known size is reported. Sheets of steel mesh and glass capillaries were used as systems of rigid pores. The roots of grasses were found to be capable of penetrating pores much smaller than their nominal thickness, this capability being limited by the size of the root cap and the stele. Constricted root tips elongated at a slower rate but could grow down long capillaries if adequately aerated. The size of rigid pore critical to the growth of perennial ryegrass was 315 m.     

Negative plant-soil feedbacks may limit persistence of an invasive tree due to rapid accumulation of soil pathogens

Soil organisms influence plant species coexistence and invasion potential. Plant-soil feedbacks occur when plants change soil community composition such that interactions with that soil community in turn may positively or negatively affect the performance of conspecifics. Theories predict and studies show that invasions may be promoted by stronger negative soil feedbacks for native compared with exotic species. We present a counter-example of a successful invader with strong negative soil feedbacks apparently caused by host-specific, pathogenic soil fungi. Using a feedback experiment in pots, we investigated whether the relative strength of plant-soil feedbacks experienced by a non-native woody invader, Sapium sebiferum, differed from several native tree species by examining their performance in soils collected near conspecifics ('home soils') or heterospecifics ('away soils') in the introduced range. Sapium seedlings, but no native seedlings, had lower survival and biomass in its home soils compared with soils of other species (negative feedback'). To investigate biotic agents potentially responsible for the observed negative feedbacks, we conducted two additional experiments designed to eliminate different soil taxa ('rescue experiments'). We found that soil sterilization (pot experiment ) or soil fungicide applications (pot and field experiments) restored Sapium performance in home soil thereby eliminating the negative feedbacks we observed in the original experiment. Such negative feedbacks apparently mediated by soil fungi could have important effects on persistence of this invader by limiting Sapium seedling success in Sapium dominated forests (home soils) though their weak effects in heterospecific (away) soils suggest a weak role in limiting initial establishment.     

Differential response to frequency-dependent interactions: an experimental test using genotypes of an invasive grass

Positive feedbacks have been suggested as a means for non-indigenous species to successfully invade novel environments. Frequency-dependent feedbacks refer to a species performance being dependent on its local abundance in the population; however, frequency dependence is often described as a monolithic trait of a species rather than examining the variation in response for individual genotypes and fitness traits. Here, we investigate frequency-dependent outcomes for individual genotypes and fitness-related traits for the invasive grass Phalaris arundinacea. We tested for competition-mediated frequency dependence by establishing hexagonal arrays with the center target plant surrounded by either same, different or no genotype neighbors to determine how changing the small-scale frequency neighborhood-influenced invasion success. We used a Bayesian ANOVA approach which allowed us to easily accommodate our non-normal dataset and found that same neighbor plots had greater biomass production than different neighbor plots. Target plants also had greater stem height and aboveground biomass when surrounded by same genotype neighbors. A greenhouse experiment did not support the hypothesis that increased mycorrhizal associations were the cause of positive frequency dependence. We devised a frequency-dependent metric to quantify the extent of fitness-related differences for individual genotypes and found that individual genotypes showed a range of both positive and negative responses to different frequency treatments; however, only positive responses were statistically significant. The small-scale genotypic neighborhood had no effect for the fitness-related traits of leaf number, belowground biomass and total biomass. We demonstrate that individual invasive genotypes respond differently to changing frequency neighborhoods and that growth responses do not respond with the same direction and magnitude. A range of frequency-dependent responses may allow genotypes to invade a wide range of environments.     

Contrasting trait responses to latitudinal climate variation in two lineages of an invasive grass

Plants are expected to respond to global environmental change through shifts in functional traits and in their ranges. These shifts could alter productivity and interactions among species or genetic lineages, ultimately leading to changes in distributions and abundance. In particular, cosmopolitan species are predicted to increase growth with decreasing latitude due to differences in climate and temperature. The pattern of changes in growth may vary among genotypes within species, leading to different responses with latitude. To evaluate whether climate can affect geographically distinct genotypes of cosmopolitan invasive species differently, we evaluated the trait responses of two lineages of the common reed, Phragmites australis, to variation in environmental conditions spanning North America’s Atlantic coast. Using three reciprocal transplant common gardens, we tested for the effects of garden location and plant lineage on traits related to biomass production, flowering frequency, leaf morphology, and leaf-level physiology. We found that aboveground biomass, stem density, and flowering frequency responded non-linearly to increasing latitude in one or both lineages. These results suggest that measures of plant traits over narrow latitudinal ranges may not accurately reflect organismal-level responses to global change at broad spatial scales. Given the responses to latitude that we observed in P. australis, we propose that feedbacks between growth and reproductive rate will influence range shifts in these two lineages. Such range shifts could lead to genetic admixtures, subsequently yielding more productive, locally-adapted genotypes.     

The effect of hydraulic lift on organic matter decomposition,soil nitrogen cycling,and nitrogen acquisition by a grass species

Hydraulic lift (HL) is the passive movement of water through plant roots, driven by gradients in water potential. The greater soil–water availability resulting from HL may in principle lead to higher plant nutrient uptake, but the evidence for this hypothesis is not universally supported by current experiments. We grew a grass species common in North America in two-layer pots with three treatments: (1) the lower layer watered, the upper one unwatered (HL), (2) both layers watered (W), and (3) the lower layer watered, the upper one unwatered, but with continuous light 24 h a day to limit HL (no-HL). We inserted ingrowth cores filled with enriched-nitrogen organic matter (¹⁵N-OM) in the upper layer and tested whether decomposition, mineralization and uptake of ¹⁵N were higher in plants performing HL than in plants without HL. Soils in the upper layer were significantly wetter in the HL treatment than in the no-HL treatment. Decomposition rates were similar in the W and HL treatments and lower in no-HL. On average, the concentration of NH₄ ⁺-N in ingrowth cores was highest in the W treatment, and NO₃ ⁻-N concentrations were highest in the no-HL treatment, with HL having intermediate values for both, suggesting differential mineralization of organic N among treatments. Aboveground biomass, leaf ¹⁵N contents and the ¹⁵N uptake in aboveground tissues were higher in W and HL than in no-HL, indicating higher nutrient uptake and improved N status of plants performing HL. However, there were no differences in total root nitrogen content or ¹⁵N uptake by roots, indicating that HL affected plant allocation of acquired N to photosynthetic tissues. Our evidence for the role of HL in organic matter decomposition and nutrient cycling suggests that HL could have positive effects on plant nutrient dynamics and nutrient turnover.     

Effects of exotic invasive trees on nitrogen cycling: a case study in Central Spain

We assess the hypothesis that rates of nitrogen transformations in the soil are altered upon replacement of native by exotic trees, differing in litter properties. Ailanthus altissima and Robinia pseudoacacia, two common exotic trees naturalized in the Iberian Peninsula, were compared with the native trees Ulmus minor and Fraxinus angustifolia, respectively. Naturally senesced leaves of each species were collected and C:N ratio, N and lignin content assessed. We prepared 64 litter bags per species and left them to decompose, below the canopy of the same species and below the canopy of the paired species. Dry mass, N concentration and N pool of the remaining litter were assessed after 5 and 7 months. Soil samples were collected three times during the experiment to assess soil moisture, organic matter, pH, potential mineralization rates and mineral N pools. Mineral N availability was assessed three times in the field by using ion-exchange resin-impregnated membranes. Ailanthus litter decomposed faster than Ulmus litter, probably due to the higher toughness of the latter. In spite of its high N content, Robinia litter decomposed slower than Fraxinus one, probably due to its high lignin content. In both cases, litter decomposition was faster below the exotic than the native canopies. The release of N per unit of initial litter mass was higher under both invaded situations (Ailanthus below Ailanthus and Robinia below Robinia) than under the native ones. However, soils collected below native and exotic trees neither differed in potential N mineralization rate nor in mineral N. This may be attributed to a quick plant uptake of released N and/or to a high organic matter accumulation in the soil previous to invasion that can exert a tighter control on soil N transformations than the current exotic litter.     

Variation for phenotypic plasticity among populations of an invasive exotic grass

Phenotypic plasticity is a common feature of plant invaders, but little is known about variation in plasticity among invading populations. Variation in plasticity of ecologically important traits could facilitate the evolution of greater plasticity and invasiveness. We examined plasticity among invasive populations of Microstegium vimineum (Japanese stiltgrass), a widespread and often dominant grass of forests in the eastern U.S. with two separate experiments. First, we exposed seven Microstegium populations to a drought treatment in growth chambers and monitored growth and physiological responses. Then, we established a greenhouse experiment using a subset of the populations; two that exhibited the most divergent responses and one intermediate population. In the greenhouse, we manipulated drought and shade and evaluated biomass production and specific leaf area (SLA). Microstegium exhibited plasticity for biomass production and SLA in the greenhouse experiment, and populations significantly varied in the degree of plasticity under drought and shade treatments. Two populations significantly increased biomass production under favorable conditions, unlike the third population. The most productive populations also responded to shade stress via greater SLA, possibly allowing for greater utilization of available light, while the third population did not. These results show that Microstegium can exhibit plastic responses to environmental conditions. Moreover, variation for plasticity among populations provides the potential for further evolution of plasticity. Future studies should focus on the relative importance of plasticity for the success of Microstegium and other plant invaders and evaluate post-introduction evolution of plasticity.     

The evolution of nitrogen cycling

Based upon arguments concerning properties of the environment and the energetics of nitrogen transformation reactions, new hypotheses regarding their evolution are presented. These hypotheses are supported by new calculations and observations germane to understanding the evolution of the nitrogen cycle. From calculations of shock production by meteor impact, we suggest that impact produced fixed nitrogen could have resulted in the entire reservoir of Earth's N₂ being converted into fixed nitrogen at the end of accretion. We have significantly improved upon previous calculations of the abiotic fixation rate on the early earth and find a rate of fixation by lightning of 1–3 × 10¹⁶ Molecules NO/J, which is 2 to 3 times greater than previous estimates. This strengthens the suggestion, corroborated by the predominance of a single nitrogenase enzyme, that biological nitrogen fixation may have been a late evolutionary development, after the development of an aerobic atmosphere. In addition, we show for the first time that HNO, predicted to be the main product of atmospheric photochemical reactions involving NO on the primitive Earth by photochemical models, would eventually become NO₂ ^– and NO₃ ^– after reaching the Earth's surface. Based upon microbe-environment interactions on an ecological as well as a biochemical scale we suggest that denitrification arose prior to aerobic respiration and that nitrification arose after the advent of an aerobic atmosphere. We hypothesize the following evolutionary sequence for the biological transformation of nitrogen compounds: Ammonification Denitrification Nitrification Nitrogen fixation.