The effects of a 9-year nitrogen and water addition on soil aggregate phosphorus and sulfur availability in a semi-arid grassland

Previous studies have demonstrated that higher nitrogen (N) and water availability affect both above- and below-ground communities, soil carbon and N pools, and microbial activity in semi-arid grasslands of Inner Mongolia. However, how soil phosphorus (P) and sulfur (S) pools, and related soil enzyme activities (as indicators of P and S cycles) respond to long-term N and water addition has still remained unclear. Since 2005, a field experiment with urea and water amendments has been conducted to examine their effects on total and available P and S concentrations and alkaline phosphomonoesterase (PME) and aryl-sulfatase (ArS) activities in three soil aggregate fractions: large macroaggregates (>2 mm), small macroaggregates (0.25–2 mm), and microaggregates (<0.25 mm) in an Inner Mongolia semi-arid grassland. Normalized to aggregate mass, microaggregates retained the highest total P and S concentrations. Both N and water additions increased the available P (by up to 84.5%) and the available S (by up to 150%) in the soil aggregate fractions. Soil acidification, as a result of the N addition, decreased both alkaline PME and ArS activities by up to 62.9% and 39.6%, respectively, while the water addition increased their activities. Our observations revealed that soil acidification (under the N addition) and elevated enzyme activity (under the water addition) played important roles in the levels of soil available P and S. The depression of P- and S-acquiring enzymes with soil acidification may decrease P and S availability, potentially impacting ecosystem processes and limiting the restoration of these grassland systems. The water addition was shown to be a more effective practice than the urea amendment for improving soil structure, supplying available P and S, and maintaining the sustainability of this semi-arid grassland.     

Increased competition does not lead to increased phylogenetic overdispersion in a native grassland

That competition is stronger among closely related species and leads to phylogenetic overdispersion is a common assumption in community ecology. However, tests of this assumption are rare and field‐based experiments lacking. We tested the relationship between competition, the degree of relatedness, and overdispersion among plants experimentally and using a field survey in a native grassland. Relatedness did not affect competition, nor was competition associated with phylogenetic overdispersion. Further, there was only weak evidence for increased overdispersion at spatial scales where plants are likely to compete. These results challenge traditional theory, but are consistent with recent theories regarding the mechanisms of plant competition and its potential effect on phylogenetic structure. We suggest that specific conditions related to the form of competition and trait conservatism must be met for competition to cause phylogenetic overdispersion. Consequently, overdispersion as a result of competition is likely to be rare in natural communities.     

Resource competition and suppression of plants colonizing early successional old fields

Early colonizing annual plants are rapidly suppressed in secondary succession on fertile midwestern old fields, while later colonizing perennials persist. Differences in competitive ability for above- and belowground resources may be partly responsible for differences in species persistence during succession, as both light and nutrient availability may change rapidly. We found that, although both above- and belowground competition suppress growth of colonizing plants, belowground competition was the dominant factor in the suppression of the annual Ambrosia artemisiifolia in 2nd-year-old fields near the W.K. Kellogg Biological Station in southwestern Michigan. Despite an ability to persist in later successional fields, seedling transplants of the perennial Achillea millefolium were also suppressed by above- and belowground competition, with belowground competition having the strongest effect. As in many old fields, nitrogen availability is the primary factor limiting plant productivity. There was no clear difference between the species in ability to compete for ¹⁵N from an enriched patch, although there was an indication of greater precision of foraging by Achillea. Life history differences between these species and consequent differences in the phenology of root growth relative to other old-field plants are likely to play a large role in the persistence of Achillea in successional fields where Ambrosia is suppressed. Received: 8 January 1998 / Accepted: 16 September 1998     

Nutrient resorption response to fire and nitrogen addition in a semi-arid grassland

Fire and nitrogen (N) addition, both widely used grassland restoration strategies, strongly influence community composition and ecosystem functioning. However, little is known about their effects on plant nutrient resorption from senescing leaves, especially in semi-arid ecosystems. We evaluated the effects of fire, N addition (5.25 g N m⁻² yr⁻¹) and their potential interactions on nutrient resorption in five plant species in a semi-arid grassland in northern China. Foliar nutrient concentrations and resorption proficiencies and efficiencies varied substantially among species and functional groups. Fire increased green leaf N concentration ([N]g) and decreased N resorption proficiency (N RP), P resorption proficiency (P RP) and P resorption efficiency (P RE). N addition led to higher [N]g and lower N resorption, whereas it did not affect P related responses. There was no interaction between fire and N addition to affect all response variables except for green leaf P concentration ([P]g). These results suggest that fire and N addition can influence ecosystem nutrient cycling directly by changing resorption patterns and litter quality. Given the substantial interspecific variations in nutrient content and resorption and the potentially changing community composition, both fire and N addition may have indirect impacts on ecosystem nutrient cycling in this semi-arid grassland.     

施肥与竞争生长对入侵植物黄顶菊和本地植物光合特性的影响

【背景】黄顶菊具有较强的生态适应性和竞争能力,可压制或排挤本地物种,形成单优势种群,进而导致当地生物多样性下降。【方法】选取黄顶菊入侵域的4种伴生植物益母草、苘麻、马唐、反枝苋为研究对象,采用LI-6400便携式光合仪测定入侵植物黄顶菊与这4种植物竞争生长过程中的光合特性,并研究施肥对不同植物与黄顶菊竞争生长过程中光合速率、蒸腾速率和胞间CO2浓度的影响。【结果】竞争生长对黄顶菊净光合速率的影响与本地植物种类密切相关,施肥促进了黄顶菊的净光合速率;竞争生长显著抑制了益母草、马唐和反枝苋的净光合速率,促进了苘麻的净光合速率,施肥增强了这4种植物的净光合速率。竞争生长对益母草和苘麻蒸腾速率的影响显著,而对其他2种植物的影响较小;与苘麻、益母草和反枝苋竞争生长过程中黄顶菊的蒸腾速率显著降低;施肥显著降低了单种处理中黄顶菊的蒸腾速率。与益母草、苘麻和反枝苋竞争生长过程中黄顶菊的胞间CO2浓度明显降低,施肥处理对黄顶菊的CO2利用率有显著抑制作用。【结论与意义】施肥影响黄顶菊的光合特性,与不同本地植物竞争生长对黄顶菊光合特性的影响不同。本研究结果将为进一步揭示黄顶菊的入侵机制及其综合治理提供理论依据。     

Microbial responses to nitrogen addition in three contrasting grassland ecosystems

The effects of global N enrichment on soil processes in grassland ecosystems have received relatively little study. We assessed microbial community response to experimental increases in N availability by measuring extracellular enzyme activity (EEA) in soils from three grasslands with contrasting edaphic and climatic characteristics: a semiarid grassland at the Sevilleta National Wildlife Refuge, New Mexico, USA (SEV), and mesic grasslands at Konza Prairie, Kansas, USA (KNZ) and Ukulinga Research Farm, KwaZulu-Natal, South Africa (SAF). We hypothesized that, with N enrichment, soil microbial communities would increase C and P acquisition activity, decrease N acquisition activity, and reduce oxidative enzyme production (leading to recalcitrant soil organic matter [SOM] accumulation), and that the magnitude of response would decrease with soil age (due to higher stabilization of enzyme pools and P limitation of response). Cellulolytic activities followed the pattern predicted, increasing 35–52% in the youngest soil (SEV), 10–14% in the intermediate soil (KNZ) and remaining constant in the oldest soil (SAF). The magnitude of phosphatase response did not vary among sites. N acquisition activity response was driven by the enzyme closest to its pH optimum in each soil: i.e., leucine aminopeptidase in alkaline soil, β-N-acetylglucosaminidase in acidic soil. Oxidative enzyme activity varied widely across ecosystems, but did not decrease with N amendment at any site. Likewise, SOM and %C pools did not respond to N enrichment. Between-site variation in both soil properties and EEA exceeded any treatment response, and a large portion of EEA variability (leucine aminopeptidase and oxidative enzymes), 68% as shown by principal components analysis, was strongly related to soil pH (r = 0.91, P < 0.001). In these grassland ecosystems, soil microbial responses appear constrained by a molecular-scale (pH) edaphic factor, making potential breakdown rates of SOM resistant to N enrichment.     

Determinants of growing season soil CO2flux in a Minnesota grassland

Soil CO₂ flux was measured across 947 plots at 7 experimentalgrassland sites at the Cedar Creek Natural History Area in order to determinethe relationships between soil CO₂ flux and environmental factors,living plant biomass, and soil C and N. Soil CO₂ flux increased asthe day progressed, and was positively related to aboveground biomass,belowground biomass, and soil % C. However, most of the variation in soilCO₂ flux explained by a multiple regression model(r ² = 0.55) was attributed to the different experimental sites (61%).Soil CO₂ flux increased with increasing aboveground plant biomass(explaining 16% of the model variation),belowground plant biomass (12%), and soil C and C:N ratio(6%). The length of time between aboveground biomass in aplot was clipped and soil CO₂ flux variedamong plots. Soil CO₂ flux declined with increased timesince clipping, supporting the idea that recently fixedcarbon is a significant component of soil CO₂ flux.Soil CO₂ flux did not follow standard Q₁₀relationships. Over a 20 °C temperature range,soil CO₂ flux tended to be lower in warmer plots.More work is necessary to understand what factors explainthe large differences that were seen among experimentalsites in soil CO₂ flux that could not be explainedby biomass or soil properties.     

Neighbourhood-scale diversity, composition and root crowding do not alter competition during drought in a native grassland

The diversity‐invasion resistance theory argues that increased diversity results in increased competitive suppression of establishing plants. Although there is support for the pattern of decreased invasion with increased diversity, empirical demonstrations of increased competition are limited. An experiment was conducted during a severe drought in a native grassland community. The drought resulted in minimal shading among neighbours, and in contrast to prior studies, competition here was exclusively belowground. Neither diversity nor community composition influenced root crowding or competition. It appears that when competition is belowground, it is independent of diversity, likely because of fundamental differences in the mechanisms of above‐ and belowground competition. This suggests that even at the neighbourhood scale, there is no inherently negative relationship between competition and diversity, and lends support to alternative theories which suggest factors of than diversity may more strongly influence community invasibility.     

Fertilization alters nitrogen isotopes and concentrations in ectomycorrhizal fungi and soil in pine forests

To assess how nitrogen (N) availability affected ectomycorrhizal functioning and to test a theoretical model of ectomycorrhizal ¹⁵N partitioning, we measured C/N and δ¹⁵N in soils and nine fungal taxa in two Swedish N addition experiments. Sporocarp C/N and soil C/N decreased with fertilization, implying that N uptake per unit fungal growth increased. The S horizon was more responsive than the F and H horizons to changes in N addition, with N turnover for these horizons of 24, 57, and 57 y, respectively. Fungal and soil δ¹⁵N patterns identified fungal N sources, with N acquisition primarily from the S, F, or H horizon for two, five, and two taxa, respectively. With increasing N availability, sporocarp ¹⁵N enrichment increased in five taxa, in agreement with our model of fungal-plant N partitioning. However, it decreased in Lactarius rufus and Russula aeruginea, perhaps indicating shifts towards greater inorganic N uptake in these two taxa. This may relate to the generally lower sensitivity of these taxa to N deposition compared to the Cortinarius and Suillus taxa that fit our model of ¹⁵N partitioning.     

光照条件对土壤-植物系统氮素状况影响的研究

应用盆栽试验,通过调节不同光照强度并控制其它条件相互一致的条件下,研究了光照条件对土壤植物系统N素状况以及作物(莴笋)产量的影响.结果表明,光照强度的改变会引起作物生长状况的相应变化,同时也导致土壤N素(NH₄⁺-N、NO₃^--N)状况、作物吸收N量以及作物对N素吸收速度等的改变.在试验所处的光照强度范围内,光照较强时,则作物吸收N素的速度较快、吸收N量增加,且产量高,但土壤中相应的N素含量(NH₄⁺-N、NO₃^--N)则只能维持在相对较低的水平;光照较弱时,则出现与此相反的情况.     

连续氮添加14年对温带典型草原土壤碳氮组分及物理结构的影响

全球氮沉降对生态系统造成了深远的影响,研究长时间氮沉降对草地生态系统土壤理化特征的影响有助于加强生态系统对氮沉降响应的长效机制的理解。通过连续14年长期施加N0（0 g N m^-2 a^-1）、N2（2 g N m^-2 a^-1）、N4（4 g N m^-2 a^-1）、N8（8 g N m^-2 a^-1）、N16（16 g N m^-2 a^-1）、N32（32 g N m^-2 a^-1）六种浓度尿素模拟氮沉降,并将土壤分成0-10、10-20和20-40 cm三个深度土层,研究温带草原生态系统土壤碳氮组分及物理结构对氮添加的响应及其相互关系,结果表明：（1）氮添加显著降低0-10 cm土壤酸碱度及土壤微生物量碳含量,N32相比N0分别下降了27.63%和58.40%（P<0.05）;各土层总有机碳和全氮含量对氮添加处理无显著响应,0-10 cm土层显著高于20-40 cm土层。（2）同一土层深度不同梯度氮添加处理显著增加土壤无机氮离子含量（P<0.05）,0-10 cm土层铵态氮含量N32相比N0增加了88.72%,20-40 cm土层硝态氮含量N32相比N0增加了19.55倍,土壤深度与氮添加对无机氮离子含量影响具有显著的交互效应。（3）同一土壤深度不同梯度氮添加处理土壤粒度分形维数及土壤团聚体差异不显著,相关分析表明土壤碳氮元素含量与土壤结构显著相关。土壤碳氮组分在适宜浓度氮添加的增加趋势说明氮添加在一定程度上可能促进土壤理化性质的改良,氮添加对土壤物理结构的影响还需要进一步的深入研究。     

Biomass partitioning and water content relationships at the branch and whole-plant levels and as a function of plant size in Elaeagnus mollis populations in Shanxi, North China

Understanding of the biomass (dry weight) allocation and water relations in populations will provide useful information on the growth patterns and resource-allocation dynamics. By destructive sampling, foliage, branch and root biomass were measured in the endangered shrub Elaeagnus mollis populations growing in Shanxi province, North China. Biomass partitioning and water content relationships were compared at the branch and whole-plant levels, and as a function of basal diameter (plant size). The biomass was mainly distributed in the bigger branches at the branch level, and in the branch wood at the whole-plant level, and branch biomass (but not foliage or root biomass) increases significantly with increasing basal diameter. As a result, branch wood became the major biomass pool, even though considerable biomass was also allocated to the roots. However, the relative water content decreased from the periphery of the crown to the interior of the shrub at the branch level, and from the aboveground to the belowground at the whole-plant level though no significant variation among foliage, branches, and roots. Yet it increased significantly for the whole-plant with increasing basal diameter. The ratio of belowground to aboveground biomass was smaller than 1.0, even as a function of basal diameter. These growth responses indicated a strong adaptation to the shrub’s growing conditions. Biomass was primarily allocated above the ground and the aboveground components grew faster than the belowground one.     

光照条件对土壤—植物系统氮素状况影响的研究

应用盆栽试验 ,通过调节不同光照强度并控制其它条件相互一致的条件下 ,研究了光照条件对土壤 植物系统N素状况以及作物 (莴笋 )产量的影响 .结果表明 ,光照强度的改变会引起作物生长状况的相应变化 ,同时也导致土壤N素 (NH 4 N、NO-3 N)状况、作物吸收N量以及作物对N素吸收速度等的改变 .在试验所处的光照强度范围内 ,光照较强时 ,则作物吸收N素的速度较快、吸收N量增加 ,且产量高 ,但土壤中相应的N素含量(NH 4 N、NO-3 N)则只能维持在相对较低的水平 ;光照较弱时 ,则出现与此相反的情况 .     

Interspecific competition among grasshoppers and their effect on plant abundance in experimental field environments

Summary We tested whether grasshoppers in experimental field environments, i.e. cages (40×40 cm) placed on existing old field vegetation, (1) were limited in density by plant abundance and/or nitrogen content, (2) exhibited interspecific competition, and (3) altered the relative abundance of different plant species. We examined interactions among a pair of early season grasshopper species (May–June; Arphia conspersa and Pardalophora apiculata) and a late season pair (July–August; Melanoplus femur-rubrum and Melanoplus bivittatus). Each grasshopper species was placed in cages by itself and with another grasshopper species. Grasshoppers generally survived at higher density in fertilized cages and they reduced plant abundance relative to empty cages, suggesting that grasshoppers may be food limited at these densities. In unfertilized plots, early season grasshoppers preferred grasses (Schizachyrium scoparium and Poa pratensis) and favored the growth of forbs (especially Solidago spp.). However, late in summer, Melanoplus spp. preferred Solidago spp. and favored the growth of grasses.The pattern of grasshopper survivorship and plant reduction within these experimental environments provide preliminary support for some of the predictions of resource competition theory. Grasshoppers exhibited interspecific competition only if they significantly reduced plant biomass. If two species competed, a grasshopper species was eliminated only if the superior competitor, when living by itself, could reduce plant biomass to a significantly lower level than the inferior competitor. Competitors persisted only if they did not differ in their ability to reduce plant biomass or reduced the abundance of different plant species.     

Conservatism of responses to environmental change is rare under natural conditions in a native grassland

Whether or not niche conservatism is common is widely debated. Despite this uncertainty, closely related species are often assumed to be ecologically similar. This principle has led to the proposed use of phylogenetic information in forecasting species responses to environmental change. Tests of niche conservatism often focus on ‘functional traits’ and environmental tolerances, but there have been limited tests for conservatism in species’ responses to changes in the environment, especially in the field. The prevalence of functional convergence and the likelihood of functional trade-offs in a heterogeneous environment suggest that conservatism of the response niche is unlikely to be detectable under natural conditions. To test the relevance of evolutionary information in predicting ecological responses, we tested for conservatism (measured as phylogenetic signal) of grassland plant population responses to 14 treatments (e.g. light, nutrients, water, enemies, mutualists), each manipulated for 2–3 years, and 4 treatment categories (aboveground, belowground, resource, and herbivory) at a single site. Individual treatment responses showed limited evidence of conservatism, with only weak conservatism in plant responses to mycorrhizae and grazing. Aspects of the response niche were conserved among monocots both aboveground and belowground, although the pattern varied. Conservatism was limited to grazing aboveground, but belowground responses were conserved as a group, suggesting fundamental differences in how selection has led to niche conservatism in aboveground and belowground environments. Overall, our results suggest that conservatism of the response niche is not common, but is actually rare. As such, evolutionary relationships are likely to be of limited relevance for predicting species responses under field conditions, at least over the short time scales used in this study.     

Changes in nitrogen resorption traits of six temperate grassland species along a multi-level N addition gradient

Nitrogen (N) resorption from senescing leaves is an important mechanism of N conservation for terrestrial plant species, but changes in N-resorption traits over wide-range and multi-level N addition gradients have not been well characterized. Here, a 3-year N addition experiment was conducted to determine the effects of N addition on N resorption of six temperate grassland species belonging to three different life-forms: Stipa krylovii Roshev. (grass), Cleistogenes squarrosa (T.) Keng (grass), Artemisia frigida Willd. (semishrub), Melissitus ruthenica C.W.Wang (semishrub and N-fixer), Potentilla acaulis L. (forb) and Allium bidentatum Fisch.ex Prokh. (forb). Generally, N concentrations in green leaves increased asymptotically for all species. N concentrations in senescent leaves for most species (5/6) also increased asymptotically, except that the N concentration in senescent leaves of A. bidentatum was independent of N addition. N-resorption efficiency decreased with increasing N addition level only for S. krylovii and A. frigida, while no clear responses were found for other species. These results suggest that long-term N fertilization increased N uptake and decreased N-resorption proficiency, but the effects on N-resorption efficiency were species-specific for different temperate grassland species in northern China. These inter-specific differences in N resorption may influence the positive feedback between species dominance and N availability and thus soil N cycling in the grassland ecosystem in this region.