Short-term effects of nitrogen enrichment,litter removal and cutting on a Mediterranean grassland

Semi-natural grassland communities are of great interest in conservation because of their high species richness. These communities are being threatened by both land abandonment and nitrogen eutrophication, and their continued existence will depend upon correct management. However, there is a distinct lack of studies of the ecological mechanisms that regulate species diversity and productivity in Mediterranean grasslands. We have conducted a 3-year field experiment in a species-poor grassland in central Italy to investigate the effects of nitrogen fertilization coupled with removal of plant litter and artificial cutting on species diversity and community productivity. Vegetation cutting reduced living biomass but increased species diversity. In fact, cutting had positive effects on the cover of almost all of the annual and biennial species, while it had a negative effect on the dominant perennial grasses Brachypodium rupestre and Dactylis glomerata. Litter removal had similar effects to cutting, although it was far less effective in increasing species diversity. In contrast, nitrogen enrichment strongly increased the living biomass while maintaining very low species diversity. Our results have indicated that semi-natural Mediterranean grasslands need specific management regimes for maintenance and restoration of species diversity. In the management of these grasslands, attention should be paid to the potential threat from nitrogen enrichment, especially when coupled with land abandonment.     

Year-round cattle and horse grazing supports the restoration of abandoned,dry sandy grassland and heathland communities by supressing Calamagrostis epigejos and enhancing species richness

The spread of competitive grasses, changes in species composition and vegetation structure are direct consequences of grassland and heathland abandonment. As an alternative to more costly management measures such as traditional pastoralism, year-round low-intensity grazing with large herbivores is increasingly used to restore and maintain semi-open habitats. However, the suitability of this grazing regime has not yet been investigated for long-abandoned, highly degraded but nutrient-poor sandy grassland and heathland communities. In particular, it is unclear if year-round grazing is suitable for preventing the further spread of highly competitive grasses such as Calamagrostis epigejos while simultaneously maintaining or improving characteristic species richness and vegetation structure. Hence, we conducted a comprehensive field study on two spatial scales (plot-level: 25 m², macroplot-level: 1 ha) to analyse the impacts of year-round low-intensity cattle and horse grazing on the development of the highly competitive grass Calamagrostis epigejos, as well as the vegetation structure and plant species richness of long-abandoned but nutrient-poor dry sandy grassland and heathland communities, their mosaics and Calamagrostis stands within an 800 ha heathland between 2008 and 2015. Finally, we assessed the local conservation status of the habitat types after seven years of grazing in comparison to long-abandoned sites.Grazing successfully reduced the coverage of Calamagrostis epigejos, whereby Calamagrostis stands developed towards species-rich sandy grasslands after seven years of grazing. In addition, the quality of the vegetation structure was improved by enhancing the proportion of bare soil, while litter and grass cover, litter thickness and height of the field layer as well as the coverage of ruderal indicators were significantly reduced on grazed sites in comparison to ungrazed sites in 2015. Moreover, we found an overall positive grazing effect on species richness: Total species number, number of target species as well as subordinated target species significantly increased within the vegetation types over time.Thus, year-round low-intensity cattle and horse grazing is a suitable management tool for restoring, maintaining and even improving long-abandoned, nutrient-poor sandy grassland and heathland communities, and thus to enhance the local conservation status of the habitat types. However, if there is a high initial cover of woody species (e.g. shrubs, tree rejuvenation), then an extensive shrub and tree clearance will be necessary, with manual shrub cutting being crucial to reduce the heavy regrowth of the woody species. In addition, a one-time mowing should be implemented in highly degraded heaths to facilitate the vegetative rejuvenation of degenerate stands of Calluna vulgaris, thus improving its attractiveness for the grazing animals.     

Nitrogen deposition causes widespread loss of species richness in British habitats

We use national scale data to test the hypothesis that nitrogen (N) deposition is strongly negatively correlated with plant species richness in a wide range of ecosystem types. Vegetation plots from a national ecological surveillance programme were drawn from heathland, acid, calcareous and mesotrophic grassland habitats. Mean species number and mean plant traits were calculated for each plot and related to atmospheric N deposition. There was a significant reduction in species richness with N deposition in acid grassland and heathland even after fitting covarying factors. In acid grassland and heathland, evidence from trait changes suggested that acidification rather than increased fertility was responsible for species loss. In contrast, calcareous grassland showed evidence of eutrophication in response to increasing N deposition. Loss of species richness from chronic N deposition is apparent in infertile grasslands and heathland. Mechanisms associated with loss of species richness differ between habitats so mitigation of N deposition should be targeted to habitat type.     

Coastal heathland succession influences butterfly community composition and threatens endangered butterfly species

Succession has a strong influence on species diversity and composition of semi-natural open terrestrial ecosystems. While several studies examined the effects of succession on butterflies in grassland and forest ecosystems, the response of heathland butterflies to succession had not been investigated so far. To address this issue we sampled butterfly abundance and environmental parameters on the Baltic island of Hiddensee (NE Germany) along a gradient of coastal heathland succession from grey dunes to birch forest. Our results provide evidence that succession of coastal heathland has a strong influence on butterfly diversity, abundance, and species composition. Thereby grass and tree encroachment present the main threats for heathland butterflies. Diversity and abundance of butterflies were highest in shrub-encroached heath directly followed by early stages of coastal heathland succession (dwarf-shrub heath, grey dune). Both observed threatened species (Hipparchia semele, Plebeius argus) were negatively affected by succession: abundance decreased with increasing vegetation density (both species) and grass cover (P. argus); consequently, the two later successional stages (shrub, birch forest) were not occupied. Our findings highlight the importance of the preservation of early stages of coastal heathland succession for endangered butterfly species. For coastal heathland management we therefore suggest to maintain early successional stages by sheep grazing, mowing or, in case of high nutrient contents, intensive techniques such as sod-cutting or choppering. To a lower extend shrub-encroached sites should also be present, which might be beneficial for overall species richness.     

Above- and below-ground resource acquisition strategies determine plant species responses to nitrogen enrichment

Background and AimsKnowledge 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.MethodsBased 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.Key ResultsWe 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.ConclusionsOur 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.     

The influence of savanna trees on nutrient,water and light availability and the understorey vegetation

In an East African savanna herbaceous layer productivity and species composition were studied around Acacia tortilis trees of three different age classes, as well as around dead trees and in open grassland patches. The effects of trees on nutrient, light and water availability were measured to obtain an insight into which resources determine changes in productivity and composition of the herbaceous layer. Soil nutrient availability increased with tree age and size and was lowest in open grassland and highest under dead trees. The lower N:P ratios of grasses from open grassland compared to grasses from under trees suggested that productivity in open grassland was limited by nitrogen, while under trees the limiting nutrient was probably P. N:P ratios of grasses growing under bushes and small trees were intermediate between large trees and open grassland indicating that the understorey of Acacia trees seemed to change gradually from a N-limited to a P-limited vegetation. Soil moisture contents were lower under than those outside of canopies of large Acacia trees suggesting that water competition between trees and grasses was important. Species composition of the herbaceous layer under Acacia trees was completely different from the vegetation in open grassland. Also the vegetation under bushes of Acacia tortilis was different from both open grassland and the understorey of large trees. The main factor causing differences in species composition was probably nutrient availability because species compositions were similar for stands of similar soil nutrient concentrations even when light and water availability was different. Changes in species composition did not result in differences in above-ground biomass, which was remarkably similar under different sized trees and in open grassland. The only exception was around dead trees where herbaceous plant production was 60% higher than under living trees. The results suggest that herbaceous layer productivity did not increase under trees by a higher soil nutrient availability, probably because grass production was limited by competition for water. This was consistent with the high plant production around dead trees because when trees die, water competition disappears but the high soil nutrient availability remains. Hence, in addition to tree soil nutrient enrichment, below-ground competition for water appears to be an important process regulating tree-grass interactions in semi-arid savanna.     

Arthropod fauna on grassland–heathland associations under different grazing managements with domestic ruminants

The effects of two grazer species (cattle or sheep) and two flock types (single or mixed with goats) on vegetation and arthropod fauna were studied in a factorial design on eight plots which comprised two thirds of mechanically cleared heathland and one third of improved ryegrass-clover grassland. After six grazing seasons, the shrubland areas were dominated by gorse (Ulex gallii) in all treatments. Herbaceous cover was higher under mixed than under single grazing, and under sheep than under cattle grazing. Higher captures of Opiliones, Julida, Lithobiomorpha, Microcoryphia and Carabidae were recorded in shrublands than in grasslands, while the reverse was observed for Linyphiidae, Lycosidae and Hemiptera. Within shrublands, fauna responded to the flock type but not to the grazer species. More arthropod groups favoured the patchier areas with higher herbaceous biomass generated by mixed herds with goats. Within grasslands, species-specific responses to the grazer species were observed. Mixed grazing schemes which include goats within partially improved heathlands could contribute to maintain higher biodiversity levels in these marginal areas.     

氮水添加对放牧背景下荒漠草原生产力的影响

水分与氮素作为干旱和半干旱草原生产力的共同限制性因子在退化草原的生态快速修复过程中备受关注。以不同放牧强度背景下的短花针茅荒漠草原为研究对象,开展围封模拟放牧利用实验,同时添加氮素和水分。通过分析历史放牧强度与年份对生产力的影响,以及添加氮素和水分对不同功能群植物生物量的作用,探讨放牧强度对短花针茅草原生产力的内在作用机制,以及如何实现荒漠草原资源合理开发和可持续利用。研究结果显示,降雨量与放牧强度决定着短花针茅草原的植物群落结构。氮素和水分添加可分别提升11%-29%和12%-32%的群落地上生物量,且二者存在显著的交互作用。不同功能群植物的地上生物量对氮素与水分添加的响应存在差异,多年生丛生禾草对氮素和水分添加响应最敏感。氮素与水分添加可显著提高多年生丛生禾草的地上生物量,但与自然降水量相关。氮素添加对地上生物量的影响在正常降雨和稍旱年份作用显著,而水分添加在干旱年份作用显著。在正常降雨年份,以半灌木植物为优势种的轻度放牧背景以添加水分对提升生产力最宜,以多年生丛生禾草和半灌木为共优种的中度放牧背景和以多年生丛生禾草为优势种的重度放牧以同时添加水分和氮素对提升生产力最为宜;在干旱年份不同放牧强度背景下均以同时添加水分和氮素对提升生产力最为宜。我们的结果表明了养分与资源的改善有利于退化短花针茅草原的快速恢复和可持续生产。     

Early regeneration of Calluna heathland under various fertilization treatments

Summary A sod-cutting and fertilization experiment was performed on a Calluna-dominated heathland in The Netherlands to determine appropriate management regimes for Calluna regeneration, and to further understand the nutrient responses of heathland species. Replicated permanent plots were analysed by multivariate techniques. Sod-cutting alone caused Calluna regeneration from its soil seed bank. A single fertilization at the start of the experiment caused initial vegetation differences which disappeared after a few years as the nutrients were lost from the system, except that one application of nitrogen enhanced the rate of Calluna regeneration. Repeated fertilization caused large differences in the vegetation: repeated nitrogen enhanced several bryophyte species while greatly inhibiting Calluna, repeated phosphate partly inhibited Calluna while greatly favouring several lichen species, and the most striking result of repeated calcium was also an increase in bryophytes, but the species were different from those favoured by nitrogen. Treatments which inhibited Calluna tended to increase species diversity because of the lessened Calluna dominance.     

Assessing the impact of land abandonment, nitrogen enrichment and fairy-ring fungi on plant diversity of Mediterranean grasslands

Semi-natural calcareous grasslands are of great conservation interest because of their high species richness, but they are threatened by land abandonment and nitrogen eutrophication. These plant communities evolved as a result of a long history of human activity, which generated and maintained these habitats by extensive grazing and mowing. Calcareous grasslands are listed as a priority for conservation in the EC Habitats Directive. However, the effects of different management regimes, nitrogen enrichment, and soil-borne pathogens on plant species diversity are less clear for grasslands of the Mediterranean Basin, compared to meadows in Northern and Central Europe. In this study, we assessed the impact of land abandonment, nitrogen enrichment, and fairy-ring fungi on species diversity in semi-natural grasslands found in the Mediterranean Basin by comparing the available literature with findings from recent studies carried out in Central Italy. In a series of field experiments, the cutting of abandoned grassland consistently reduced the living biomass of the dominant perennial grasses, such as Brachypodium rupestre and Bromus erectus, and promoted a rapid increase in species richness and diversity by allowing the establishment of rare species. There was a similar, but less effective, restoration of species diversity and composition in mowed grassland after litter removal. We also show that nitrogen enrichment at levels comparable to atmospheric deposition depresses species diversity, which also hampers the positive effects of litter removal. Our findings are consistent with previous results achieved in Northern and Central Europe, which however, mainly focused on grasslands with intermediate to high primary productivity levels. The limited availability of data from low-productivity, drought-prone Mediterranean grasslands requires further studies to assess the impact of land abandonment and nitrogen eutrophication in such ecosystems. Finally, we discuss the role of fairy-ring fungi in the maintenance of plant diversity in species-rich grassland. We show that fairy-ring fungi (e.g. Agaricus campestris) critically affect the spatial distribution and diversity of coexisting plant species. By killing the dominant perennial herbs, these radially growing plant pathogens produce empty niches for rare, short-lived species, thus affecting the vegetation pattern. Overall, our results are of interest for environmental managers, as they provide guidelines for the restoration of abandoned areas and the conservation of these species-rich habitats.     

Interactive effects of resource enrichment and resident diversity on invasion of native grassland by <Emphasis Type="Italic">Lolium arundinaceum</Emphasis>

Resident diversity and resource enrichment are both recognized as potentially important determinants of community invasibility, but the effects of these biotic and abiotic factors on invasions are often investigated separately, and little work has been done to directly compare their relative effects or to examine their potential interactions. Here, we evaluate the individual and interactive effects of resident diversity and resource enrichment on plant community resistance to invasion. We factorially manipulated plant diversity and the enrichment of belowground (soil nitrogen) and aboveground (light) resources in low-fertility grassland communities invaded by Lolium arundinaceum, the most abundant invasive grass in eastern North America. Soil nitrogen enrichment enhanced L. arundinaceum performance, but increased resident diversity dampened this effect of nitrogen enrichment. Increased light availability (via clipping of aboveground vegetation) had a negligible effect on community invasibility. These results demonstrate that a community’s susceptibility to invasion can be contingent upon the type of resource pulse and the diversity of resident species. In order to assess the generality of these results, future studies that test the effects of resident diversity and resource enrichment against a range of invasive species and in other environmental contexts (e.g., sites differing in soil fertility and light regimes) are needed. Such studies may help to resolve conflicting interpretations of the diversity–invasibility relationship and provide direction for management strategies.     

Short-term effects of nitrogen enrichment, litter removal and cutting on a Mediterranean grassland

Semi-natural grassland communities are of great interest in conservation because of their high species richness. These communities are being threatened by both land abandonment and nitrogen eutrophication, and their continued existence will depend upon correct management. However, there is a distinct lack of studies of the ecological mechanisms that regulate species diversity and productivity in Mediterranean grasslands. We have conducted a 3-year field experiment in a species-poor grassland in central Italy to investigate the effects of nitrogen fertilization coupled with removal of plant litter and artificial cutting on species diversity and community productivity. Vegetation cutting reduced living biomass but increased species diversity. In fact, cutting had positive effects on the cover of almost all of the annual and biennial species, while it had a negative effect on the dominant perennial grasses Brachypodium rupestre and Dactylis glomerata. Litter removal had similar effects to cutting, although it was far less effective in increasing species diversity. In contrast, nitrogen enrichment strongly increased the living biomass while maintaining very low species diversity. Our results have indicated that semi-natural Mediterranean grasslands need specific management regimes for maintenance and restoration of species diversity. In the management of these grasslands, attention should be paid to the potential threat from nitrogen enrichment, especially when coupled with land abandonment.     

Long‐term effects of artificial nighttime lighting and trophic complexity on plant biomass and foliar carbon and nitrogen in a grassland community

The introduction of artificial nighttime lighting due to human settlements and transport networks is increasingly altering the timing, intensity, and spectra of natural light regimes worldwide. Much of the research on the impacts of nighttime light pollution on organisms has focused on animal species. Little is known about the impacts of daylength extension due to outdoor lighting technologies on wild plant communities, despite the fact that plant growth and development are under photoperiodic control. In a five‐year field experiment, artificial ecosystems (“mesocosms”) of grassland communities both alone or in combination with invertebrate herbivores and predators were exposed to light treatments that simulated street lighting technologies (low‐pressure sodium, and light‐emitting diode [LED]‐based white lighting), at ground‐level illuminance. Most of the plant species in the mesocosms did not exhibit changes in biomass accumulation after 5 years of exposure to the light treatments. However, the white LED treatment had a significant negative effect on biomass production in the herbaceous species Lotus pedunculatus. Likewise, the interaction between the white LED treatment and the presence of herbivores significantly reduced the mean shoot/root ratio of the grass species Holcus lanatus. Artificial nighttime lighting had no effect on the foliar carbon or nitrogen in most of the grassland species. Nevertheless, the white LED treatment significantly increased the leaf nitrogen content in Lotus corniculatus in the presence of herbivores. Long‐term exposure to artificial light at night had no general effects on plant biomass responses in experimental grassland communities. However, species‐specific and negative effects of cool white LED lighting at ground‐level illuminance on biomass production and allocation in mixed plant communities are suggested by our findings. Further studies on the impacts of light pollution on biomass accumulation in plant communities are required as these effects could be mediated by different factors, including herbivory, competition, and soil nutrient availability.     

On the water relations of four heath species

Four species of heath, occurring in the heathlands of Brittany, are compared regarding their water relations: Calluna vulgaris, Erica ciliaris, E. cinerea and E. tetralix. E. cinerea is unable to establish itself in wet heathland because of its intolerance of prolonged waterlogging. It is the Erica species best adapted to dry habitat conditions in Brittany. E. ciliaris canot establish itself in dry heathland and is less tolerant of waterlogging than E. tetralix. E. tetralix is the species best adapted to wet heathland, being tolerant of waterlogging, but can also establish itself in dry heathland. Of the three E. species it has the widest ecological range. Calluna is tolerant of both wet and dry conditions and has a wide ecological range. E. cinerea is typical of dry- and E. tetralix of wet heathland. Although both species did best in moist aerated soil in experimental cultures, neither is abundant in mesophilic heaths where E. ciliaris is dominant. One explanation may be competition for aerial space. E. cinerea and E. tetralix both have an upright growth, whereas E. ciliaris rapidly adopts a straggling bushy habit, with long rooting branches. E. ciliaris thus establishes large interpenetrating clumps. With increasing dryness E. ciliaris disappears and may be replaced by E. cinerea and, with increasing wetness and especially waterlogging, E. tetralix will take over.     

Fire increases aboveground biomass,seed production and recruitment success of Molinia caerulea in dry heathland

During the last decades, the perennial tussock grass Molinia caerulea has shown an increased abundance in European heathlands, most likely as a result of increased nitrogen deposition and altered management schemes. Because of its deciduous nature, Molinia produces large amounts of litter each year, which may affect the intensity and frequency of accidental fires in heathlands. These fires may influence plant population dynamics and heathland community organization through their effects on plant vital attributes and competitive interactions. In this study, fire-induced changes in competitive ability and invasiveness of Molinia through changes in biomass production, seed set and seed germination under both natural and laboratory conditions were investigated. We found that fire significantly increased aboveground biomass, seed set and germination of Molinia. Seed set was twice as high in burned compared to unburned heathland. Two years after fire, seedling densities in natural conditions were on average six times higher in burned than in unburned heathland, which resulted in increased abundance of Molinia after burning. The seed germination experiment indicated that seeds harvested from plants in burned heathland showed higher germination rates than those from unburned heathland. Hence, our results clearly demonstrate increased invasive spread of Molinia after large and intense fires. Active management guidelines are required to prevent further encroachment of Molinia and to lower the probability of large fires altering the heathland community in the future.     

Nitrogen deposition and the reduction of butterfly biodiversity quality in the Netherlands

Butterfly decline in Northern Europe is a cause of concern and it has been hypothesised that this is due to nitrogen deposition inducing excess early growth of plants. It has also been changing the quality of the food available to larvae. We tested these hypotheses by linking butterfly biodiversity quality indices (species richness, population, biomass, conservation value, evenness (Simpson's Index) and modelled species richness (Chao 1 and 2)) with nitrogen Critical Load Exceedence (nCLE) data. An index of butterfly sensitivity to nitrogen was also created (Species Nitrogen Value Index (SNVI)). Using PCA, datasets were tested for associations and relationships.The results included multiple biodiversity quality indices based on 17 years of data (aggregated into three periods of six, six and five years to give 287 datasets) in four habitat types (grassland, heathland, woodland and farmland). With the exception of heathland the analysis showed that nitrogen deposition and all other indices (except SNVI) were in decline.For heathland the last 11 years did not show any significant decline. Heathland also showed an anomalous biodiversity quality profile for these last 11 years, suggesting that the sensitivity of heathland to nitrogen deposition will require further considerable efforts to achieve a nitrogen deposition that is not in exceedence of the critical load.Habitat restoration will take time due to the multiple hindrances to colonisation, which in the case of heathland might prevent successful butterfly colonisation for the foreseeable future. These results indicate the efficacy of butterfly biodiversity quality and nCLE as indicators for the SEBI 2020 process (Streamlining European Biodiversity Indicators) by showing the relationship between them.     

内蒙古温带草原大针茅叶片光合生理特性对氮添加的响应

大气氮沉降增加生态系统氮有效性,优势种植物对不同水平氮输入的响应影响草原生态系统结构和功能。研究设置4个氮添加水平,分析内蒙古温带草原优势种大针茅（Stipa grandis）光合生理特性对不同梯度氮添加的响应。结果表明：低氮（0-2 g m^-2 a^-1）处理时,大针茅叶片氮含量较低,叶绿素含量和1,5-二磷酸核酮糖羧化/加氧酶的活性不高,光能利用效率低,导致光系统II出现过剩激发能,光合器官受到抑制,净光合速率相对较低。适量氮添加（5-10 g m^-2 a^-1）提高了大针茅叶片羧化系统和电子传递系统的氮分配,进而提高了1,5-二磷酸核酮糖羧化/加氧酶的活性以及电子传递速率,净光合速率增大。高氮（25 g m^-2 a^-1）处理时,叶片氮含量较高,但光合氮分配比例下降,降低了光合氮利用效率。大针茅光抑制程度增大,叶绿素含量、1,5-二磷酸核酮糖羧化/加氧酶的活性下降,不利于生物量积累。研究结果有助于进一步了解全球变化背景下草原生态系统优势种的生理响应机制,并为草原的可持续发展提供一定的理论依据。     

Hierarchical plant responses and diversity loss after nitrogen addition: testing three functionally-based hypotheses in the Inner Mongolia grassland

Background

Numerous studies have shown that nitrogen (N) deposition decreases biodiversity in terrestrial ecosystems. To explain the N-induced species loss, three functionally based hypotheses have been proposed: the aboveground competition hypothesis, the belowground competition hypothesis, and the total competition hypothesis. However, none of them is supported sufficiently by field experiments. A main challenge to testing these hypotheses is to ascertain the role of shoot and root competition in controlling plant responses to N enrichment. Simultaneously examining both aboveground and belowground responses in natural ecosystems is logistically complex, and has rarely been done.

Methodology/Principal Findings

In a two-year N addition experiment conducted in a natural grassland ecosystem, we investigated both above- and belowground responses of plants at the individual, species, and community levels. Plants differed significantly in their responses to N addition across the different organizational levels. The community-level species loss was mainly due to the loss of perennial grasses and forbs, while the relative abundance of plant species was dependent mainly on individual-level responses. Plasticity in biomass allocation was much smaller within a species than between species, providing a biological basis for explaining the functionally based species loss. All species increased biomass allocation to aboveground parts, but species with high belowground allocations were replaced by those with high aboveground allocations, indicating that the increased aboveground competition was the key process responsible for the observed diversity loss after N addition in this grassland ecosystem.

Conclusions/Significance

Our findings shed new light on the validity of the three competing hypotheses concerning species loss in response to N enrichment. They also have important implications for predicting the future impacts of N deposition on the structure and functioning of terrestrial ecosystems. In addition, we have developed a new technique for ascertaining the roles of aboveground and belowground competition in determining plant responses to N fertilization.     

Nitrate Reduction to Nitrite,Nitric Oxide and Ammonia by Gut Bacteria under Physiological Conditions

The biological nitrogen cycle involves step-wise reduction of nitrogen oxides to ammonium salts and oxidation of ammonia back to nitrites and nitrates by plants and bacteria. Neither process has been thought to have relevance to mammalian physiology; however in recent years the salivary bacterial reduction of nitrate to nitrite has been recognized as an important metabolic conversion in humans. Several enteric bacteria have also shown the ability of catalytic reduction of nitrate to ammonia via nitrite during dissimilatory respiration; however, the importance of this pathway in bacterial species colonizing the human intestine has been little studied. We measured nitrite, nitric oxide (NO) and ammonia formation in cultures of Escherichia coli, Lactobacillus and Bifidobacterium species grown at different sodium nitrate concentrations and oxygen levels. We found that the presence of 5 mM nitrate provided a growth benefit and induced both nitrite and ammonia generation in E.coli and L.plantarum bacteria grown at oxygen concentrations compatible with the content in the gastrointestinal tract. Nitrite and ammonia accumulated in the growth medium when at least 2.5 mM nitrate was present. Time-course curves suggest that nitrate is first converted to nitrite and subsequently to ammonia. Strains of L.rhamnosus, L.acidophilus and B.longum infantis grown with nitrate produced minor changes in nitrite or ammonia levels in the cultures. However, when supplied with exogenous nitrite, NO gas was readily produced independently of added nitrate. Bacterial production of lactic acid causes medium acidification that in turn generates NO by non-enzymatic nitrite reduction. In contrast, nitrite was converted to NO by E.coli cultures even at neutral pH. We suggest that the bacterial nitrate reduction to ammonia, as well as the related NO formation in the gut, could be an important aspect of the overall mammalian nitrate/nitrite/NO metabolism and is yet another way in which the microbiome links diet and health.     

草原群落退化演替过程中微斑块土壤碳氮的空间异质动态

微斑块变化是草原退化过程中的活跃成分。分析了呼伦贝尔克氏针茅草原逆行演替过程中微斑块土壤全碳、全氮和碱解氮含量的空间异质性,提出了"养分聚集效应"的概念。研究结果表明:随着群落退化演替的加剧,土壤全碳、全氮和碱解氮的含量均表现为演替前期演替后期演替中期(P0.05)。从土壤全碳、全氮和碱解氮的变异系数和变异函数综合分析来看,10 cm×10 cm微尺度上,草原退化演替过程中土壤全碳、全氮和碱解氮的空间异质性具有明显的不一致性;全碳的空间异质性表现为演替中期演替前期演替后期,全氮表现为演替后期演替前期演替中期,碱解氮表现为演替中期演替后期演替前期。草原退化过程中土壤养分在微斑块上的富积和迁移表现出尺度依赖性和变异性。