Temporal variability of foliar nutrients: responses to nitrogen deposition and prescribed fire in a temperate steppe

Plant nutrient concentrations and stoichiometry drive fundamental ecosystem processes, with important implications for primary production, diversity, and ecosystem sustainability. While a range of evidence exists regarding how plant nutrients vary across spatial scales, our understanding of their temporal variation remains less well understood. Nevertheless, we know nutrients regulate plant function across time, and that important temporal controls could strongly interact with environmental change. Here, we report results from a 3-year assessment of inter-annual changes of foliar nitrogen (N) and phosphorus (P) concentrations and stoichiometry in three dominant grasses in response to N deposition and prescribed fire in a temperate steppe of northern China. Foliar N and P concentrations and their ratios varied greatly among years, with this temporal variation strongly related to inter-annual variation in precipitation. Nitrogen deposition significantly increased foliar N concentrations and N:P ratios in all species, while fire significantly altered foliar N and P concentrations but had no significant impacts on N:P ratios. Generally, N addition enhanced the temporal stability of foliar N and decreased that of foliar P and of N:P ratios. Our results indicate that plant nutrient status and response to environmental change are temporally dynamic and that there are differential effects on the interactions between environmental change drivers and timing for different nutrients. These responses have important implications for consideration of global change effects on plant community structure and function, management strategies, and the modeling of biogeochemical cycles under global change scenarios.     

Hierarchical responses of plant stoichiometry to phosphorus addition in an alpine meadow community

为了探明氮(N)限制的植物群落中物种水平和功能群水平的碳(C)、N、磷(P)含量以及C:N:P对P添加的响应是否一致,明确P添加对群落物种构成改变的内在机制。以青藏高原高寒草甸为研究对象, 通过P添加试验, 研究了功能群水平和物种水平生态化学计量比对P添加的响应, 以及P添加对物种水平的优势度和功能群水平生物量的影响。结果表明: 在青藏高原高寒草甸连续5年添加P显著改变了植物的C、N、P含量以及C:N:P, 且在物种水平和功能群水平(不含典型物种)的响应规律基本一致。在禾本科、莎草科和杂类草功能群(不含典型物种)和相应物种水平上P添加对C含量影响不显著。P添加显著增加了禾本科、莎草科、豆科和杂类草4个功能群(不含典型物种)和相应物种水平的植物P含量, 降低了C:P和N:P。禾本科和莎草科的N含量和C:N对P添加在物种水平和功能群水平上(不含典型物种)的响应规律一致, 表现为N含量显著降低, C:N显著增加; P添加使豆科物种水平上N含量显著增加而C:N显著降低, 但在功能群水平上(不含典型物种)无显著作用; 杂类草的N含量和C:N对P添加在物种水平和功能群水平上(不含典型物种)的响应规律均不一致。在N限制的生境中添加P, 禾本科物种在群落中逐渐占据优势跟其增高的N、P利用效率相关, 而杂类草由于逐渐降低的N和P利用效率使其生物量在群落中所占的比重逐渐下降。     

高寒草甸植物化学计量比对磷添加的分层响应

为了探明氮(N)限制的植物群落中物种水平和功能群水平的碳(C)、N、磷(P)含量以及C:N:P对P添加的响应是否一致,明确P添加对群落物种构成改变的内在机制。以青藏高原高寒草甸为研究对象, 通过P添加试验, 研究了功能群水平和物种水平生态化学计量比对P添加的响应, 以及P添加对物种水平的优势度和功能群水平生物量的影响。结果表明: 在青藏高原高寒草甸连续5年添加P显著改变了植物的C、N、P含量以及C:N:P, 且在物种水平和功能群水平(不含典型物种)的响应规律基本一致。在禾本科、莎草科和杂类草功能群(不含典型物种)和相应物种水平上P添加对C含量影响不显著。P添加显著增加了禾本科、莎草科、豆科和杂类草4个功能群(不含典型物种)和相应物种水平的植物P含量, 降低了C:P和N:P。禾本科和莎草科的N含量和C:N对P添加在物种水平和功能群水平上(不含典型物种)的响应规律一致, 表现为N含量显著降低, C:N显著增加; P添加使豆科物种水平上N含量显著增加而C:N显著降低, 但在功能群水平上(不含典型物种)无显著作用; 杂类草的N含量和C:N对P添加在物种水平和功能群水平上(不含典型物种)的响应规律均不一致。在N限制的生境中添加P, 禾本科物种在群落中逐渐占据优势跟其增高的N、P利用效率相关, 而杂类草由于逐渐降低的N和P利用效率使其生物量在群落中所占的比重逐渐下降。     

Nitrogen fertilization and fire act independently on foliar stoichiometry in a temperate steppe

Nitrogen (N) fertilization, as a grassland management strategy, has been widely used to improve forage quality and increase the productivity of grasslands degraded by overstocking. It is widely accepted that N addition will alter ecosystem structure and function, and that these effects may be altered by natural disturbances, such as fire. We examined the effects of annual burning and N fertilization (17.5 g N m^?2 year^?1, at a surplus rate in order to simulate agriculture treatment) on foliar chemistry and stoichiometric ratios of eight dominant species (Leymus chinensis, Stipa grandis, Cleistogenes squarrosa, Potentilla bifurca, Thalictrum squarrosum, Artemisia frigida, Kochia prostrata and Caragana microphylla) in a temperate steppe in northern China. After 3 years of treatments, annual burning significantly increased soil extractable phosphorus (P) concentration but showed no effects on soil inorganic N concentration, whereas N fertilization caused a significant increase in inorganic N concentration but not of extractable P. Species differed substantially with respect to all nutritional and stoichiometric variables. Both annual burning and N fertilization caused significant increases in foliar N and P concentrations and thus decreases in carbon (C):N and C:P ratios. While annual burning showed no effects on N:P ratios, N fertilization produced higher N:P ratios. However, species responded idiosyncratically to both fire and N fertilization in terms of foliar N concentration, C:N and N:P ratio. In addition, there was no interaction between fire and N fertilization that affected all variables. This study suggests that both annual burning and N fertilization have direct impacts on plant elemental composition and that fire- and N addition-induced changes of community composition may have important consequences for plant-mediated biogeochemical cycling pathways in temperate steppe ecosystem.     

Differential responses of auto‐ and heterotrophic soil respiration to water and nitrogen addition in a semiarid temperate steppe

Evaluating how autotrophic (SR_A), heterotrophic (SR_H) and total soil respiration (SR_TOT) respond differently to changes of environmental factors is critical to get an understanding of ecosystem carbon (C) cycling and its feedback processes to climate change. A field experiment was conducted to examine the responses of SR_A and SR_H to water and nitrogen (N) addition in a temperate steppe in northern China during two hydrologically contrasting growing seasons. Water addition stimulated SR_A and SR_H in both years, and their increases were significantly greater in a dry year (2007) than in a wet year (2006). N addition increased SR_A in 2006 but not in 2007, while it decreased SR_H in both years, leading to a positive response of SR_TOT in 2006 but a negative one in 2007. The different responses of SR_A and SR_H indicate that it will be uncertain to predict soil C storage if SR_TOT is used instead of SR_H to estimate variations in soil C storage. Overall, N addition is likely to enhance soil C storage, while the impacts of water addition are determined by its relative effects on carbon input (plant growth) and SR_H. Antecedent water conditions played an important role in controlling responses of SR_A, SR_H and the consequent SR_TOT to water and N addition. Our findings highlight the predominance of hydrological conditions in regulating the responses of C cycling to global change in the semiarid temperate steppe of northern China.     

Linkages of plant stoichiometry to ecosystem production and carbon fluxes with increasing nitrogen inputs in an alpine steppe

Unprecedented levels of nitrogen (N) have entered terrestrial ecosystems over the past century, which substantially influences the carbon (C) exchange between the atmosphere and biosphere. Temperature and moisture are generally regarded as the major controllers over the N effects on ecosystem C uptake and release. N‐phosphorous (P) stoichiometry regulates the growth and metabolisms of plants and soil organisms, thereby affecting many ecosystem C processes. However, it remains unclear how the N‐induced shift in the plant N:P ratio affects ecosystem production and C fluxes and its relative importance. We conducted a field manipulative experiment with eight N addition levels in a Tibetan alpine steppe and assessed the influences of N on aboveground net primary production (ANPP), gross ecosystem productivity (GEP), ecosystem respiration (ER), and net ecosystem exchange (NEE); we used linear mixed‐effects models to further determine the relative contributions of various factors to the N‐induced changes in these parameters. Our results showed that the ANPP, GEP, ER, and NEE all exhibited nonlinear responses to increasing N additions. Further analysis demonstrated that the plant N:P ratio played a dominate role in shaping these C exchange processes. There was a positive relationship between the N‐induced changes in ANPP (ΔANPP) and the plant N:P ratio (ΔN:P), whereas the ΔGEP, ΔER, and ΔNEE exhibited quadratic correlations with the ΔN:P. In contrast, soil temperature and moisture were only secondary predictors for the changes in ecosystem production and C fluxes along the N addition gradient. These findings highlight the importance of plant N:P ratio in regulating ecosystem C exchange, which is crucial for improving our understanding of C cycles under the scenarios of global N enrichment.     

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。     

Water-mediated responses of ecosystem carbon fluxes to climatic change in a temperate steppe

Global warming and a changing precipitation regime could have a profound impact on ecosystem carbon fluxes, especially in arid and semiarid grasslands where water is limited. A field experiment manipulating temperature and precipitation has been conducted in a temperate steppe in northern China since 2005. A paired, nested experimental design was used, with increased precipitation as the primary factor and warming simulated by infrared radiators as the secondary factor. The results for the first 2 yr showed that gross ecosystem productivity (GEP) was higher than ecosystem respiration, leading to net C sink (measured by net ecosystem CO(2) exchange, NEE) over the growing season in the study site. The interannual variation of NEE resulted from the difference in mean annual precipitation. Experimental warming reduced GEP and NEE, whereas increased precipitation stimulated ecosystem C and water fluxes in both years. Increased precipitation also alleviated the negative effect of experimental warming on NEE. The results demonstrate that water availability plays a dominant role in regulating ecosystem C and water fluxes and their responses to climatic change in the temperate steppe of northern China.     

Testing nitrogen and water co-limitation of primary productivity in a temperate steppe

Background and aims

Studies have found significant differences in methane (CH₄) emissions among rice cultivars; however, it is unclear whether this difference is related to radial oxygen loss (ROL) from the roots.

Methods

Based on a 2-year in situ field study and solution culture experiments on 16 rice cultivars, we investigated CH₄ emission levels and their dependence on ROL.

Results

We detected significant differences in CH₄ emission and ROL among rice cultivars. The lowest and highest CH₄ emission levels were 4.10 and 7.35 g m^?2 for early rice, and 14.36 and 23.33 g m^?2 for late rice, respectively. The maximum and minimum ROL values were 3.77 and 1.73 mmol plant^?1 h^?1 for early rice, and 4.18 and 2.08 mmol plant^?1 h^?1 for late rice, respectively. Seasonal total CH₄ emission was negatively correlated with ROL in the early rice season (p?<?0.01), and (p?<?0.01) in the late rice season. ROL was positively correlated with the number of roots per plant (RN), root tips per plant (RT), and root volume per plant (RV).

Conclusions

We suggest that ROL can be used as a predictive index for CH₄ emissions. RN, RT, and RV were the most important factors influencing ROL in rice cultivars.

     

Effects of increased nitrogen deposition and precipitation on seed and seedling production of Potentilla tanacetifolia in a temperate steppe ecosystem

Background

The responses of plant seeds and seedlings to changing atmospheric nitrogen (N) deposition and precipitation regimes determine plant population dynamics and community composition under global change.

Methodology/Principal Findings

In a temperate steppe in northern China, seeds of P. tanacetifolia were collected from a field-based experiment with N addition and increased precipitation to measure changes in their traits (production, mass, germination). Seedlings germinated from those seeds were grown in a greenhouse to examine the effects of improved N and water availability in maternal and offspring environments on seedling growth. Maternal N-addition stimulated seed production, but it suppressed seed mass, germination rate and seedling biomass of P. tanacetifolia. Maternal N-addition also enhanced responses of seedlings to N and water addition in the offspring environment. Maternal increased-precipitation stimulated seed production, but it had no effect on seed mass and germination rate. Maternal increased-precipitation enhanced seedling growth when grown under similar conditions, whereas seedling responses to offspring N- and water-addition were suppressed by maternal increased-precipitation. Both offspring N-addition and increased-precipitation stimulated growth of seedlings germinated from seeds collected from the maternal control environment without either N or water addition. Our observations indicate that both maternal and offspring environments can influence seedling growth of P. tanacetifolia with consequent impacts on the future population dynamics of this species in the study area.

Conclusion/Significance

The findings highlight the importance of the maternal effects on seed and seedling production as well as responses of offspring to changing environmental drivers in mechanistic understanding and projecting of plant population dynamics under global change.     

Root trait responses of six temperate grassland species to intensive mowing and NPK fertilisation: a field study in a temperate grassland

Background and aims

Plant traits may characterize functional ecosystem properties and help to predict community responses to environmental change. Since most traits used relate to aboveground plant organs we aim to explore the indicative value of root traits.

Methods

We examined the response of root traits (specific root length [SRL], specific root surface area [SRA], root diameter [RD], root tissue mass density [TMD], root N concentration) in six grassland species (3 grasses, 3 herbs) to four management regimes (low vs. high mowing frequency; no fertilization vs. high NPK fertilization). The replicated experiment in temperate grassland with long continuity simulated the increase in grassland management intensity in the past 50 years in Central Europe.

Results

Increasing mowing frequency (one vs. three cuts per year) led to no significant root trait changes. NPK fertilization resulted in considerable trait shifts with all species responding in the same direction (higher SRL, SRA and N concentration, lower TMD) but at different magnitude. Fertilization-driven increases in SRA were mainly caused by lowered tissue density while root diameter reduction was the main driver of SRL increases.

Conclusion

We conclude that root morphological traits may be used as valuable indicators of environmental change and increasing fertilization in grasslands.     

Soil microbial properties and plant growth responses to carbon and water addition in a temperate steppe: the importance of nutrient availability

Background

Global climatic change is generally expected to stimulate net primary production, and consequently increase soil carbon (C) input. The enhanced C input together with potentially increased precipitation may affect soil microbial processes and plant growth.

Methodology/Principal Findings

To examine the effects of C and water additions on soil microbial properties and plant growth, we conducted an experiment lasting two years in a temperate steppe of northeastern China. We found that soil C and water additions significantly affected microbial properties and stimulated plant growth. Carbon addition significantly increased soil microbial biomass and activity but had a limited effect on microbial community structure. Water addition significantly increased soil microbial activity in the first year but the response to water decreased in the second year. The water-induced changes of microbial activity could be ascribed to decreased soil nitrogen (N) availability and to the shift in soil microbial community structure. However, no water effect on soil microbial activity was visible under C addition during the two years, likely because C addition alleviated nutrient limitation of soil microbes. In addition, C and water additions interacted to affect plant functional group composition. Water addition significantly increased the ratio of grass to forb biomass in C addition plots but showed only minor effects under ambient C levels. Our results suggest that soil microbial activity and plant growth are limited by nutrient (C and N) and water availability, and highlight the importance of nutrient availability in modulating the responses of soil microbes and plants to potentially increased precipitation in the temperate steppe.

Conclusions/Significance

Increased soil C input and precipitation would show significant effects on soil microbial properties and plant growth in the temperate steppe. These findings will improve our understanding of the responses of soil microbes and plants to the indirect and direct climate change effects.     

Effects of nitrogen addition and mowing on rodent damage in an Inner Mongolian steppe

Rodent damage is a serious threat to sustainable management of grassland. The effects of nitrogen (N) deposition and grassland management on rodent damage have been scarcely studied. Here, we reported the effects of 2 years of N addition and mowing on burrow density and damage area of Citellus dauricus in a semiarid steppe in Inner Mongolia. N addition significantly aggravated, while mowing alleviated rodent damage in the grassland under study. Burrow density and damage area increased 2.8‐fold and 4.7‐fold, in N addition plots compared to the ambient N addition treatment, respectively. Conversely, mowing decreased burrow density and damage area by 75.9% and 14.5%, respectively, compared to no mowing plots. Observed changes in rodent damage were mainly due to variations in plant community cover, height, and aboveground net primary productivity. Our findings demonstrate that N addition and mowing can affect the rodent density and activity in grassland, suggesting that the effects of a changing atmospheric composition and land use on rodent damage must be considered in order to achieve better grassland management.     

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...     

Warming changes plant competitive hierarchy in a temperate steppe in northern China

Aims Quantifying changes in plant growth and interspecific interactions, both of which can alter dominance of plant species, will facilitate explanation and projection of the shifts in species composition and community structure in terrestrial biomes expected under global warming. We used an experimental warming treatment to examine the potential influence of global warming on plant growth and interspecific interactions in a temperate steppe in northern China.Materials and Methods Six dominant plant species were grown in monoculture and all 15 two-species mixtures for one growing season under ambient and elevated temperatures in the field. Temperature was manipulated with infrared radiators.Important findings Total biomass of all the six plant species was increased by 34–63% in monocultures and 20–76% in mixtures. The magnitude of the warming effect on biomass was modified by plant interactions. Experimental warming changed the hierarchies of both competitive response and competitive effect. The competitive ability (in terms of response and effect) of one C₄ grass (Pennisetum centrasiaticum) was suppressed, while the competitive abilities of one C₃ forb (Artemisia capillaris) and one C₃ grass (Stipa krylovii) were enhanced by experimental warming. The demonstrated alterations in growth and plant interactions may lead to changes in community structure and biodiversity in the temperate steppe in a warmer world in the future.     

典型草原建群种羊草对氮磷添加的生理生态响应

由于人类活动和气候变化的共同作用, 大气氮(N)沉降日益加剧, 使得陆地生态系统中的可利用性N显著增加, 生态系统更易受其他元素如磷(P)的限制。然而, 目前关于N、P养分添加对草原生态系统不同组织水平的影响研究较少, 相关机制尚不清楚。该文以内蒙古典型羊草(Leymus chinensis)草原为研究对象, 通过连续两年(2011-2012年)的N和P养分添加实验, 研究建群种羊草的生理生态性状、种群生物量和群落初级生产力对N、P添加的响应及其适应机制。结果表明: 羊草草原不同组织水平对N、P添加的响应不同。群落水平上, 地上净初级生产力在不同降水年份均受N和P元素的共同限制, N、P共同添加显著提高了地上净初级生产力; 物种水平上, N、P添加对羊草种群生物量和密度, 以及相对生物量均没有显著影响, 表明羊草能够维持种群的相对稳定; 个体水平上, 在正常降水年份(2011年), 羊草生长主要受N素限制, 而在湿润年份(2012年), 降水增加使得羊草生长没有受到明显的养分限制。羊草通过增加比叶面积、叶片大小和叶片N含量, 提高整体光合能力, 以促进个体生长。总之, 内蒙古典型草原群落净初级生产力受N、P元素共同限制, 作为建群种的羊草, 其对N、P添加的响应因组织水平而异, 也受年际间降水变化的影响。     

Soil invertebrate and plant responses to mowing and carbofuran application in a North American tallgrass prairie

Relationships between soil invertebrate populations and primary production of a tallgrass prairie were investigated using the insecticide-nematicide carbofuran and a range of mowing intensities to manipulate invertebrate densities and resource quantity and quality. The trophic composition of nematode populations was monitored through each of two growing seasons. Earthworm and macroarthropod densities and primary production were assessed at the end of the second season. Invertebrate densities were generally reduced in carbofuran-treated plots, although individual weights of surviving macroarth-ropod herbivores increased significantly (p<0.05). Carbofuran failed to affect estimates of above- or belowground plant biomass after two years of treatment. Changes in resource quantity and quality resulted in rapid responses by dominant invertebrate consumer populations. A 28% reduction in live root mass and a 24% increase in root detritus following two years of mowing was associated with a 54% decrease in herbivorous nematode densities, a 47% increase in microbivorous nematode densities, and a 41% increase in native earthworm biomass.     

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.     

Response of bacterial communities and plant-mediated soil processes to nitrogen deposition and precipitation in a desert steppe

Plant and Soil - Changes in nitrogen (N) and precipitation levels can substantially alter soil properties and plant growth, thereby altering soil microbial diversity and functionality. We used...