Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption‐mediated, positive plant–soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.     

A global comparison of grassland biomass responses to CO2 and nitrogen enrichment

Grassland ecosystems cover vast areas of the Earth''s surface and provide many ecosystem services including carbon (C) storage, biodiversity preservation and the production of livestock forage. Predicting the future delivery of these services is difficult, because widespread changes in atmospheric CO₂ concentration, climate and nitrogen (N) inputs are expected. We compiled published data from global change driver manipulation experiments and combined these with climate data to assess grassland biomass responses to CO₂ and N enrichment across a range of climates. CO₂ and N enrichment generally increased aboveground biomass (AGB) but effects of CO₂ enrichment were weaker than those of N. The response to N was also dependent on the amount of N added and rainfall, with a greater response in high precipitation regions. No relationship between response to CO₂ and climate was detected within our dataset, thus suggesting that other site characteristics, e.g. soils and plant community composition, are more important regulators of grassland responses to CO₂. A statistical model of AGB response to N was used in conjunction with projected N deposition data to estimate changes to future biomass stocks. This highlighted several potential hotspots (e.g. in some regions of China and India) of grassland AGB gain. Possible benefits for C sequestration and forage production in these regions may be offset by declines in plant biodiversity caused by these biomass gains, thus necessitating careful management if ecosystem service delivery is to be maximized. An approach such as ours, in which meta-analysis is combined with global scale model outputs to make large-scale predictions, may complement the results of dynamic global vegetation models, thus allowing us to form better predictions of biosphere responses to environmental change.     

Plant community responses to increased precipitation and belowground litter addition: Evidence from a 5‐year semiarid grassland experiment

Global climate change is predicted to stimulate primary production and consequently increases litter inputs. Changing precipitation regimes together with enhanced litter inputs may affect plant community composition and structure, with consequent influence on diversity and ecosystem functioning. Responses of plant community to increased precipitation and belowground litter addition were examined lasting 5 years in a semiarid temperate grassland of northeastern China. Increased precipitation enhanced community species richness and abundance of annuals by 16.8% and 44%, but litter addition suppressed them by 25% and 54.5% after 5 years, respectively. During the study period, perennial rhizome grasses and forbs had consistent negative relationship under ambient plots, whereas positive relationship between the two functional groups was found under litter addition plots after 5 years. In addition, increased precipitation and litter addition showed significant interaction on community composition, because litter addition significantly increased biomass and abundance of rhizome grasses under increased precipitation plots but had no effect under ambient precipitation levels. Our findings emphasize the importance of water availability in modulating the responses of plants community to potentially enhanced litter inputs in the semiarid temperate grassland.     

Plant community responses to nitrogen addition and increased precipitation: the importance of water availability and species traits

Global nitrogen (N) enrichment and changing precipitation regimes are likely to alter plant community structure and composition, with consequent influences on biodiversity and ecosystem functioning. Responses of plant community structure and composition to N addition and increased precipitation were examined in a temperate steppe in northern China. Increased precipitation and N addition stimulated and suppressed community species richness, respectively, across 6 years (2005–2010) of the manipulative experiment. N addition and increased precipitation significantly altered plant community structure and composition at functional groups levels. The significant relationship between species richness and soil moisture (SM) suggests that plant community structure is mediated by water under changing environmental conditions. In addition, plant height played an important role in affecting the responses of plant communities to N addition, and the effects of increased precipitation on plant community were dependent on species rooting depth. Our results highlight the importance and complexity of both abiotic (SM) and biotic factors (species traits) in structuring plant community under changing environmental scenarios. These findings indicate that knowledge of species traits can contribute to mechanistic understanding and projection of vegetation dynamics in response to future environmental change.     

水氮添加对内蒙古温带典型草原植被氮含量季节和年际动态的影响

作为对全球变化响应最敏感的生态系统类型之一,草原生态系统植被氮含量的季节、年际变化及其对气候变化(氮沉降、降水格局改变)的响应研究相对匮乏。基于内蒙古温带典型草原5年的氮添加(10 g N m^-2 a^-1)和水添加(添加量80 mm,分2 mm×40次、5 mm×16次、10 mm×8次、20 mm×4次、40 mm×2次5种处理)控制试验分析了水氮添加后植被氮含量在生态系统和物种两个水平的季节和年际变化。结果表明,高强度水添加处理有降低(10 mm/次和40 mm/次)生态系统氮含量的趋势,但不显著,小强度水添加处理(2 mm/次、5 mm/次)在不同年份之间无一致的升高或降低趋势,但所有水添加处理有降低两种优势物种整个生长季氮含量的趋势。氮添加促进生态系统和两种优势物种整个生长季的氮含量,但该促进作用可被水添加抵消,且这种抵消作用随水氮添加年限的延长而加剧。水氮添加均增加了生态系统氮含量的年际变异,但对特定物种季节内变异的影响在湿润和干旱年份存在一定差异。本研究将为预测草原生态系统对未来氮沉降增加和降水格局改变的响应及模型改进提供科学依据...     

Ecosystem responses to water and nitrogen amendment in a California grassland

The world's ecosystems are experiencing simultaneous changes in the supply of multiple limiting resources. Two of these, water and nitrogen (N) can strongly limit grassland production and can affect community composition and biogeochemical cycles in different ways. Grassland ecosystems in California may be particularly vulnerable to current and predicted changes in precipitation and N deposition, and ecosystem responses to potential interactive effects of water and N are not well understood. Here, we show strong colimitation of plant production resulting from factorial addition of water and N. In addition, water and N addition in combination led to increased dominance of the two most abundant grass species, while N addition regardless of water availability led to decreased species diversity. Late season carbon (C) flux response to water addition depended on N. Only plots that received additional water, but not N, still showed net ecosystem C uptake at the end of the experiment. Our results suggest that grassland ecosystem response to N deposition will be strongly dependent on future precipitation patterns.     

Molecular approach for assessing responses of microbial-feeding nematodes to burning and chronic nitrogen enrichment in a native grassland

A substantial proportion of the primary productivity in grassland ecosystems is allocated belowground, sustaining an abundant and diverse community of microbes and soil invertebrates. These belowground communities drive many important ecosystem functions and are responsive to a variety of environmental changes. Nematodes, an abundant and diverse component of grassland soil communities, are particularly responsive to altered environmental conditions, such as those associated with reduced fire frequency and nitrogen enrichment, with the most consistent responses displayed by microbial-feeding nematodes. However, much of the available research characterizing nematode responses to environmental change has been carried out at the taxonomic level of family or by broad trophic categories (e.g. fungivores, bacterivores). The extent to which differential responses to environmental change occurs at the genus level or below is unclear. Therefore, the objective of this study was to use molecular methods to quantify the response of microbial-feeding nematodes, at the lowest levels of taxonomic resolution, to nitrogen enrichment and changes in fire frequency. Using sequencing and quantitative polymerase chain reaction (PCR) probes for the 18S ribosomal RNA gene and the ITS1 region, we identified 19 microbial-feeding nematode taxa across four families. When nematodes were sampled across treatments, we found that some nematode taxa within a family responded similarly to nitrogen and burning treatments, while other taxa within the same family respond quite differently. Additionally, although nematodes from different families on average responded differently to nitrogen enrichment and burning, similar responses were seen in nematode taxa that span three taxonomic families. Thus, if nematodes are to be used as indicators of environmental change, care should be taken to assess the response at the lowest taxonomic level possible.     

Predominant role of water in regulating soil and microbial respiration and their responses to climate change in a semiarid grassland

Climate change can profoundly impact carbon (C) cycling of terrestrial ecosystems. A field experiment was conducted to examine responses of total soil and microbial respiration, and microbial biomass to experimental warming and increased precipitation in a semiarid temperate steppe in northern China since April 2005. We measured soil respiration twice a month over the growing seasons, soil microbial biomass C (MBC) and N (MBN), microbial respiration (MR) once a year in the middle growing season from 2005 to 2007. The results showed that interannual variations in soil respiration, MR, and microbial biomass were positively related to interannual fluctuations in precipitation. Laboratory incubation with a soil moisture gradient revealed a constraint of the temperature responses of MR by low soil moisture contents. Across the 3 years, experimental warming decreased soil moisture, and consequently caused significant reductions in total and microbial respiration, and microbial biomass, suggesting stronger negatively indirect effects through warming‐induced water stress than the positively direct effects of elevated temperature. Increased evapotranspiration under experimental warming could have reduced soil water availability below a stress threshold, thus leading to suppression of plant growth, root and microbial activities. Increased precipitation significantly stimulated total soil and microbial respiration and all other microbial parameters and the positive precipitation effects increased over time. Our results suggest that soil water availability is more important than temperature in regulating soil and microbial respiratory processes, microbial biomass and their responses to climate change in the semiarid temperate steppe. Experimental warming caused greater reductions in soil respiration than in gross ecosystem productivity (GEP). In contrast, increased precipitation stimulated GEP more than soil respiration. Our observations suggest that climate warming may cause net C losses, whereas increased precipitation may lead to net C gains in the semiarid temperate steppe. Our findings highlight that unless there is concurrent increase in precipitation, the temperate steppe in the arid and semiarid regions of northern China may act as a net C source under climate warming.     

Mycorrhizal fungi reduce the negative effects of nitrogen enrichment on plant community structure in dune grassland

Nitrogen (N) inputs to ecosystems have increased worldwide, often leading to large changes in plant community structure and reducing plant diversity. Yet, the interaction of increased N availability with other factors that determine plant community composition, are still poorly understood. Here, we test whether the impact of N addition on plant communities depends on the presence of arbuscular mycorrhizal fungi (AMF). AMF are widespread plant symbionts that facilitate growth of many plant species. We hypothesize that AM fungi reduce the negative impact of N addition on plant communities by supporting growth of species that are sensitive to N enrichment.We established experimental grassland microcosms consisting of 18 plant species. These microcosms were subjected to high and low N supply and were inoculated with AMF or remained nonmycorrhizal. Both N addition and AMF had a big impact on plant community composition, but with opposite effects. N addition induced a 2.8‐fold increase in grass biomass and reduced legume biomass. Grasses dominated the microcosms at high N supply, especially when AMF were absent. In contrast, AMF enhanced biomass of all legumes species (on average 6.8‐fold) and reduced the relative abundance of grasses. The proportion of legume biomass out of total shoot biomass at high N supply was 19% with AMF and only 3% without AMF. Our results show that responses of plant communities to N enrichment depend on AMF and that AMF can reduce the negative impact of increased N availability on plant community structure by reducing grass dominance.     

长期施肥和增水对半干旱草地土壤性质和植物性状的影响

本文对内蒙古多伦退化草地2005年建立的长期野外控制试验中施肥和增水对土壤性质和植物群落特征的影响进行了总结和综合评述.结果表明:加氮导致了表土酸化并降低酸缓冲容量,提高了表土中碳氮磷硫有效性及DTPA-浸提态铁锰铜含量,导致盐基离子钙镁钾钠总量的消耗,降低了土壤微生物群落多样性,促进了优势植物物种叶片对氮磷硫钾及锰铜锌的吸收,抑制植物叶片对铁的吸收,而对钙镁吸收无显著影响,增加了植物地上净初级生产力(ANPP),降低了植物物种多样性和群落稳定性.单独加磷增加了表土全磷和Olsen-P含量及真菌丰度,促进了植物叶片对氮、磷、硫的吸收,但对其他土壤基本化学性质及ANPP、物种多样性无显著影响.增水提高了植物群落对干旱的抵抗力,但对ANPP增长的贡献受到土壤氮有效性的限制.增水对于加氮导致的土壤酸化、植物和微生物多样性降低等具有一定的缓冲作用;加氮增水和加磷增水下,土壤微生物多样性及功能受地上植物群落结构及功能变化的影响.长期野外控制试验对于深入理解草地生态系统结构和功能对环境变化的响应具有重要意义,但单点的研究结果仍需与不同区域多点控制试验的联网研究相结合,深入开展地上与地下生态过程的关联研究,才能深入理解草地生态系统生态学的相关机制.     

Indirect effect of nitrogen enrichment modified invertebrate herbivory through altering plant community composition in an alpine meadow

Aims Nitrogen enrichment may affect ‘community invertebrate herbivory’ (hereafter ‘herbivory’) directly by changing plant species’ specific herbivory, or indirectly by altering the composition of natural plant communities. Here, we investigated how community composition altered the community herbivory in natural ecosystems and compared the relative importance of direct and indirect effects of nitrogen addition on community herbivory.     

荒漠草原植物群落结构及其稳定性对增水和增氮的响应

通过在荒漠草原开展增水和增氮野外控制试验,研究增水和增氮对荒漠草原植物群落结构、物种多样性及群落稳定性的影响。结果表明：（1）增水和增氮处理显著影响了荒漠草原植物群落结构和地上生物量,而对植物群落稳定性影响不显著（P>0.05）。增水处理显著增加了豆科和禾本科植物地上生物量（101.3%和57.9%）（P<0.05）;增水+增氮处理显著增加了植物群落盖度（43.2%）和地上生物量（112.4%）及不同功能群（禾本科和杂类草）植物盖度（75.5%和47.3%）和地上生物量（139.3%和85.7%）（P<0.05）。与增氮处理相比,增水+增氮处理显著增加了植物群落和不同功能群（禾本科和杂类草）植物高度、盖度和地上生物量（P<0.05）。（2）增水、增氮和增水+增氮处理均显著降低了植物群落Pielou指数（11.7%、8.7%和10.2%）（P<0.05）。（3）增水和增水+增氮处理提高了荒漠草原植物群落稳定性,而增氮处理降低了荒漠草原植物群落稳定性。增水处理荒漠草原植物群落稳定性效应大于增水+增氮处理。研究表明,荒漠草原植物群落结构受到氮沉降和降水增加的共同影响。增加降水对荒漠草原植物群落稳定性的积极效应可能会抵消部分氮沉降的消极影响,荒漠草原植物群落地上生物量及群落稳定性可能有所增加。     

Asynchronous responses of soil microbial community and understory plant community to simulated nitrogen deposition in a subtropical forest

Atmospheric nitrogen (N) deposition greatly affects ecosystem processes and properties. However, few studies have simultaneously examined the responses of both the above- and belowground communities to N deposition. Here, we investigated the effects of 8 years of simulated N deposition on soil microbial communities and plant diversity in a subtropical forest. The quantities of experimental N added (g of N m⁻² year⁻¹) and treatment codes were 0 (N0, control), 6 (N1), 12 (N2), and 24 (N3). Phospholipid fatty acids (PLFAs) analysis was used to characterize the soil microbial community while plant diversity and coverage were determined in the permanent field plots. Microbial abundance was reduced by the N3 treatment, and plant species richness and coverage were reduced by both N2 and N3 treatments. Declines in plant species richness were associated with decreased abundance of arbuscular mycorrhizal fungi, increased bacterial stress index, and reduced soil pH. The plasticity of soil microbial community would be more related to the different responses among treatments when compared with plant community. These results indicate that long-term N deposition has greater effects on the understory plant community than on the soil microbial community and different conservation strategies should be considered.     

施氮频率和强度对内蒙古温带草原土壤pH及碳、氮、磷含量的影响

以内蒙古温带典型草原为研究对象,通过 9 个氮沉降梯度以及一年 12 次和一年 2 次的施氮频率比对,经过4年的氮沉降模拟,研究不同氮沉降强度和频率对不同土层土壤 pH 及碳、氮、磷含量的影响.结果表明: 随着施氮强度的增加,土壤 pH 及全磷含量逐渐降低,但土壤有效氮和有效磷的含量呈增加趋势,可溶性有机碳(DOC)含量无明显变化规律,而土壤全碳、全氮含量基本保持不变.与更接近于自然氮沉降状况的一年 12 次施氮相比,一年 2 次施氮明显高估了氮沉降在降低土壤 pH和增加有效磷含量上的作用,低估了氮沉降在增加土壤有效氮含量上的作用,其中0～5 cm土层差异显著.     

Shifts of growing‐season precipitation peaks decrease soil respiration in a semiarid grassland

Changing precipitation regimes could have profound influences on carbon (C) cycle in the biosphere. However, how soil C release from terrestrial ecosystems responds to changing seasonal distribution of precipitation remains unclear. A field experiment was conducted for 4 years (2013–2016) to examine the effects of altered precipitation distributions in the growing season on soil respiration in a temperate steppe in the Mongolian Plateau. Over the 4 years, both advanced and delayed precipitation peaks suppressed soil respiration, and the reductions mainly occurred in August. The decreased soil respiration could be primarily attributable to water stress and subsequently limited plant growth (community cover and belowground net primary productivity) and soil microbial activities in the middle growing season, suggesting that precipitation amount in the middle growing season is more important than that in the early, late, or whole growing seasons in regulating soil C release in grasslands. The observations of the additive effects of advanced and delayed precipitation peaks indicate semiarid grasslands will release less C through soil respiratory processes under the projected seasonal redistribution of precipitation in the future. Our findings highlight the potential role of intra‐annual redistribution of precipitation in regulating ecosystem C cycling in arid and semiarid regions.     

丛枝菌根真菌形态结构、物种多样性和群落组成对氮沉降响应研究进展

工业革命以来,人类活动输入到生态系统中的氮迅速增加,已突破地球所能承受的氮循环阈值。过量氮沉降会造成生物多样性丧失等一系列危害,严重影响生态系统结构和功能。丛枝菌根(AM)真菌能够与大约70%-80%的陆地植物种类形成共生关系,在宿主植物养分吸收、抵抗外界不良环境压力、群落动态和物种共存、生物地球化学循环等方面具有重要的作用。探究AM真菌对氮沉降的响应对认识和把握菌根真菌缓解氮沉降的负面后果,维持生态系统的结构和功能具有重要意义。本文综述了AM真菌的形态结构、物种多样性和群落组成等对氮沉降的响应机制。前人研究表明氮沉降通常降低AM真菌的根系定殖率,减少根外菌丝密度和土壤孢子密度,改变菌丝生长的时间动态;降低AM真菌多样性,改变AM真菌群落组成。氮沉降主要通过缓解植物氮限制、降低植物对菌根的依赖性、减少植物对菌根的碳分配、改变根系和土壤中菌根生物量比率、在植物根内维持稳定的菌根真菌组成作为应对未来扰动的“保险”、改变土壤资源有效性及土壤酸度等直接和间接途径影响AM真菌结构和功能。我们建议在未来研究中整合多组学手段、开展学科交叉,聚焦复杂的生物互作体系对氮沉降的响应机制,以及AM真菌对氮沉降响应的生态后果。     

Effects of the frequency and the rate of N enrichment on community structure in a temperate grassland

Aims Nitrogen (N) enrichment caused by human activities threatens biodiversity and alters plant community composition and structure. It has been found that heavy and infrequent N inputs may over-estimate species extinction, but it remains unclear whether plant community structure will equally respond to frequent reactive N enriched conditions.     

氮添加对荒漠草原植物群落组成与微生物生物量生态化学计量特征的影响

大气氮(N)沉降增加加速了土壤N循环, 引起微生物生物量碳(C):N:磷(P)生态化学计量关系失衡、植物种丧失和生态系统服务功能降低等问题。开展N添加下植物群落组成与微生物生物量生态化学计量特征关系的研究, 可为深入了解N沉降增加引起植物多样性降低的机理提供新思路。该文以宁夏荒漠草原为研究对象, 探讨了N添加下植物生物量和群落多样性的变化趋势, 分析了微生物生物量C:N:P生态化学计量特征独立及其与其他土壤因子共同对植物群落组成的影响。结果表明: N添加下猪毛菜(Salsola collina)生物量呈显著增加趋势, 牛枝子(Lespedeza potaninii)生物量呈逐渐降低趋势, 其他植物种生物量亦呈降低趋势但未达到显著水平; 沿N添加梯度, Shannon-Wiener多样性指数、Simpson优势度指数和Patrick丰富度指数均呈先略有增加后逐渐降低的趋势; N添加提高了微生物生物量N含量和N:P, 降低了微生物生物量C:N; 植物群落组成与微生物生物量N含量、微生物生物量C:N、微生物生物量N:P、土壤NO₃ ^--N浓度、土壤NH₄ ⁺-N浓度以及土壤全P含量有较强的相关关系; 微生物生物量C:N:P生态化学计量特征对植物种群生物量和群落多样性变化的独立解释力较弱, 但却与其他土壤因子共同解释了较大变差, 意味着N添加下微生物生物量C:N:P生态化学计量特征对植物群落组成的影响与其他土壤因子高度相关。     

氮素添加对呼伦贝尔草甸草原植物氮钾元素含量和计量比的影响

钾(K)是植物叶片中仅次于氮(N)的第二大营养元素,在调节植物生长发育、稳态维持和胁迫响应等方面具有重要作用。以往关于氮素输入对植物养分状况影响的研究中多关注了氮磷养分,较少关注钾及其与其他元素的计量关系。本研究以内蒙古呼伦贝尔草甸草原为对象,研究氮素添加和刈割对植物功能群水平和群落水平钾含量及计量特征(N∶K)的影响,分析功能群自身养分状况和群落组成改变对群落水平养分状况变化的相对贡献。结果表明: 为期6年的氮素添加提高了所有植物功能群的N含量以及根茎禾草和豆科植物的K含量,刈割降低了根茎禾草和丛生禾草的N含量,而对所有功能群的K含量和N∶K均无显著影响。氮素添加显著提高了群落水平植物N和K含量,刈割仅增加了群落植物的N含量。氮素添加和刈割对功能群和群落水平上的植物N∶K均无显著影响。功能群本身的养分变化情况对群落水平植物养分状况的贡献大于群落组成的贡献。在对群落水平养分特征的影响方面,功能群本身养分变化的贡献与群落组成的贡献具有负的协变关系。呼伦贝尔草甸草原植物具有较高的N∶K内稳性,能够较好地调节自身的氮钾平衡,这对氮沉降背景下维持氮钾元素计量关系具有重要意义。