Foliar Nitrogen Dynamics and Nitrogen Resorption of a Sandy Shrub Salix gordejevii in Northern China

Resorption of nitrogen (N) from senescing leaves is an important conservation mechanism that allows plants to use the same N repeatedly. Seasonal variations in leaf nitrogen of mature green and senescing leaves and N resorption in Salix gordejevii Chang, a sandy shrub in northern China, were studied. Our objective was to compare N resorption of this Salix species that successfully occupy different habitats (shifting sandland, fixed sandland and lowland) with differences in soil N availability and moisture. Nitrogen concentrations in green and senescing leaves were higher in June and July. N resorption efficiency (percentage reduction of N between green and senescing leaves) was highest at shifting sandland, intermediate at fixed sandland, and lowest at lowland. There was a clear seasonal variation in N-resorption efficiency, with a lower value at the early growing season and a higher value during summer. N resorption efficiency was lower at the sites with higher soil N availability, suggesting that the efficiency of the resorption process is determined by the availability of the nutrient in the soil. Resorption from senescing leaves may play an important role in the nitrogen dynamics of sandy plants and reduce the nitrogen requirements for plant growth. We conclude that N resorption from senescing leaves in S. gordejevii was correlated to soil characteristics and higher N resorption on poor soils is a phenotypic adjustment by this species to maximize N-use at low availability.     

Non-Additive Effects of Water and Nitrogen Addition on Ecosystem Carbon Exchange in a Temperate Steppe

Changes in precipitation and nitrogen (N) deposition can influence ecosystem carbon (C) cycling and budget in terrestrial biomes, with consequent feedbacks to climate change. However, little is known about the main and interactive effects of water and N additions on net ecosystem C exchange (NEE). In a temperate steppe of northern China, a field-manipulated experiment was conducted to evaluate the responses of NEE and its components to improve N and water availability from 2005 to 2008. The results showed that both water and N additions stimulated gross ecosystem productivity (GEP), ecosystem respiration (ER), and NEE. Water addition increased GEP by 17%, ER by 24%, and NEE by 11% during the experimental period, whereas N addition increased GEP by 17%, ER by 16%, and NEE by 19%. The main effects of both water and N additions changed with time, with the strongest water stimulation in the dry year and a diminishing N stimulation over time. When water and N were added in combination, there were non-additive effects of water and N on ecosystem C fluxes, which could be explained by the changes in species composition and the shifts of limiting resources from belowground (water or N) to aboveground (light). The positive water and N additions effects indicate that increasing precipitation and N deposition in the future will favor C sequestration in the temperate steppe. The non-additive effects of water and N on ecosystem C fluxes suggest that multifactor experiments are better able to capture complex interactive processes, thus improving model simulations and projections.     

Effects of Multi-nutrient Additions on GHG Fluxes in a Temperate Grassland of Northern China

Human activities have substantially enhanced the availability of important nutrient elements such as nitrogen (N), phosphorus (P), and potassium (K) in ecosystems worldwide. However, how the concurrent increase in all of these nutrients will affect greenhouse gas (that is, CO₂, N₂O, CH₄) levels remains unknown. In a temperate steppe of northern China, a 2-year field experiment was conducted to examine the effects of multi-nutrient additions on GHG fluxes from 2009 to 2010. Four levels of annual nutrient loads were mimicked: 0 g NPK (control), 15.5 g P m^?2 and 19.5 g K m^?2 as KH₂PO₄ (PK), 10 g N m^?2 as NH₄NO₃ plus PK (10N + PK), and 20 g N m^?2 plus PK (20N + PK) per year. The results show that multi-nutrient additions led to significant increases in net primary production (NPP) and soil temperature (ST), a significant decrease in soil moisture (SM) in 2010, and no significant changes in other soil parameters. Seasonal patterns differed greatly for different GHG fluxes in response to different nutrient treatments, largely as a result of differences in influential factors. The 10N + PK treatment significantly increased CO₂ uptake, whereas the 20N + PK treatment significantly decreased CO₂ uptake. The application of P and K without additional N significantly enhanced CH₄ uptake, whereas the two N + PK treatments significantly enhanced N₂O emissions. Significant positive, linear relationships were found between cumulative CO₂ uptake and soil total nitrogen (TN), microbial biomass carbon, and microbial biomass nitrogen, whereas significant negative, linear relationships were found with NPP, SM, and the C/N ratio. Significant positive, linear relationships were found between cumulative N₂O emission and ST, TN, NPP, and total organic carbon, whereas no relationships were found between cumulative CH₄ uptake and any soil parameters. CO₂ flux was related to N₂O flux temporally, to a certain extent, for all the treatments. In the control, N₂O flux showed a negative, linear relationship with CH₄ flux, whereas no regular relationships were detected between CO₂ and CH₄ fluxes in any treatment. Our findings imply that increasing nutrient deposition will change the magnitude, patterns, and relationships among GHG uptakes and emissions in the future.     

Warming and Nitrogen Addition Increase Litter Decomposition in a Temperate Meadow Ecosystem

Background

Litter decomposition greatly influences soil structure, nutrient content and carbon sequestration, but how litter decomposition is affected by climate change is still not well understood.

Methodology/Principal Findings

A field experiment with increased temperature and nitrogen (N) addition was established in April 2007 to examine the effects of experimental warming, N addition and their interaction on litter decomposition in a temperate meadow steppe in northeastern China. Warming, N addition and warming plus N addition reduced the residual mass of L. chinensis litter by 3.78%, 7.51% and 4.53%, respectively, in 2008 and 2009, and by 4.73%, 24.08% and 16.1%, respectively, in 2010. Warming, N addition and warming plus N addition had no effect on the decomposition of P. communis litter in 2008 or 2009, but reduced the residual litter mass by 5.58%, 15.53% and 5.17%, respectively, in 2010. Warming and N addition reduced the cellulose percentage of L. chinensis and P. communis, specifically in 2010. The lignin percentage of L. chinensis and P. communis was reduced by warming but increased by N addition. The C, N and P contents of L. chinensis and P. communis litter increased with time. Warming and N addition reduced the C content and C:N ratios of L. chinensisand P. communis litter, but increased the N and P contents. Significant interactive effects of warming and N addition on litter decomposition were observed (P<0.01).

Conclusion/Significance

The litter decomposition rate was highly correlated with soil temperature, soil water content and litter quality. Warming and N addition significantly impacted the litter decomposition rate in the Songnen meadow ecosystem, and the effects of warming and N addition on litter decomposition were also influenced by the quality of litter. These results highlight how climate change could alter grassland ecosystem carbon, nitrogen and phosphorus contents in soil by influencing litter decomposition.     

Age-Related Modulation of the Nitrogen Resorption Efficiency Response to Growth Requirements and Soil Nitrogen Availability in a Temperate Pine Plantation

Nitrogen (N) resorption is a key strategy for conserving N in forests, and is often affected by soil nutrient condition and N sink strength within the plant. However, our understanding of the age-related pattern of N resorption and how increasing N deposition will affect this pattern is limited. Here, we investigated N resorption along a chronosequence of stands ranging in age from 2 to 100 years old, and conducted a 4-year exogenous N input experiment in stands at age class 11, 20, and 45 in a Larix Principis-rupprechtii plantation in north China. We found a logarithmic increase in leaf N resorption efficiency (NRE) and green leaf N concentration, and a logarithmic decrease in senesced-leaf N concentration along the stand-age chronosequence. Leaf NRE was negatively correlated with plant-available N concentration. Stand-level N resorption was positively correlated with the annual N requirement for tree growth. N resorption contributed to 45, 62, and 68% of the annual N supply in the 11-, 20-, and 45-year-old stands, respectively. Our exogenous N input experiment showed that leaf NRE in the 11- and 20-year-old stands decreased 17 and 12% following a 50-kg N ha^?1 y^?1 input. However, leaf NRE was not affected in the 45-year-old stand. The increases in leaf NRE and the contribution of N resorption to annual N supply along stand ages suggested that, with stand development, tree growth depends more on N resorption to supply its N need. Furthermore, the leaf NRE of mature stand was not decreased under exogenous N input, suggesting that mature stands can be stronger sinks for N deposition than young stands due to their higher capacity to retain the deposited N within plants via internal cycle. Ignoring age-related N use strategies can lead to a bias in N cycle models when evaluating forest net primary production under increasing global N deposition.     

Precipitation Pattern Regulates Soil Carbon Flux Responses to Nitrogen Addition in a Temperate Forest

Ecosystems - Changes in precipitation frequency and intensity are predicted to be more intense and frequent accompanying climate change and may have immediate or potentially prolonged effects on...     

Influence of Earthworm Invasion on Redistribution and Retention of Soil Carbon and Nitrogen in Northern Temperate Forests

We analyzed soil organic matter distribution and soil solution chemistry in plots with and without earthworms at two sugar maple (Acer saccharum)–dominated forests in New York State, USA, with differing land-use histories to assess the influence of earthworm invasion on the retention or loss of soil carbon (C) and nitrogen (N) in northern temperate forests. Our objectives were to assess the influence of exotic earthworm invasion on (a) the amount and depth distribution of soil C and N, (b) soil ¹³C and ¹⁵N, and (c) soil solution chemistry and leaching of C and N in forests with different land-use histories. At a relatively undisturbed forest site (Arnot Forest), earthworms eliminated the thick forest floor, decreased soil C storage in the upper 12 cm by 28%, and reduced soil C:N ratios from 19.2 to 15.3. At a previously cultivated forest site with little forest floor (Tompkins Farm), earthworms did not influence the storage of soil C or N or soil C:N ratios. Earthworms altered the stable isotopic signature of soil at Arnot Forest but not at Tompkins Farm; the alteration of stable isotopes indicated that earthworms significantly increased the loss of forest floor C but not N from the soil profile at Arnot Forest. Nitrate (NO₃^–) concentrations in tension and zero-tension lysimeters were much greater at Tompkins Farm than Arnot Forest, and earthworms increased NO₃^– leaching at Tompkins Farm. The results suggest that the effect of earthworm invasion on the distribution, retention, and solution chemistry of soil C and N in northern temperate forests may depend on the initial quantity and quality of soil organic matter at invaded sites.     

Enhancement of Carbon Sequestration in Soil in the Temperature Grasslands of Northern China by Addition of Nitrogen and Phosphorus

Increased nitrogen (N) deposition is common worldwide. Questions of where, how, and if reactive N-input influences soil carbon (C) sequestration in terrestrial ecosystems are of great concern. To explore the potential for soil C sequestration in steppe region under N and phosphorus (P) addition, we conducted a field experiment between 2006 and 2012 in the temperate grasslands of northern China. The experiment examined 6 levels of N (0–56 g N m^-2 yr^-1), 6 levels of P (0–12.4 g P m^-2 yr^-1), and a control scenario. Our results showed that addition of both N and P enhanced soil total C storage in grasslands due to significant increases of C input from litter and roots. Compared with control plots, soil organic carbon (SOC) in the 0–100 cm soil layer varied quadratically, from 156.8 to 1352.9 g C m^-2 with N addition gradient (R² = 0.99, P < 0.001); and logarithmically, from 293.6 to 788.6 g C m^-2 with P addition gradient (R² = 0.56, P = 0.087). Soil inorganic carbon (SIC) decreased quadratically with N addition. The net C sequestration on grassland (including plant, roots, SIC, and SOC) increased linearly from -128.6 to 729.0 g C m^-2 under N addition (R² = 0.72, P = 0.023); and increased logarithmically, from 248.5 to 698 g C m^-2under P addition (R² = 0.82, P = 0.014). Our study implies that N addition has complex effects on soil carbon dynamics, and future studies of soil C sequestration on grasslands should include evaluations of both SOC and SIC under various scenarios.     

Warming and Nitrogen Addition Alter Photosynthetic Pigments,Sugars and Nutrients in a Temperate Meadow Ecosystem

Global warming and nitrogen (N) deposition have an important influence on terrestrial ecosystems; however, the influence of warming and N deposition on plant photosynthetic products and nutrient cycling in plants is not well understood. We examined the effects of 3 years of warming and N addition on the plant photosynthetic products, foliar chemistry and stoichiometric ratios of two dominant species, i.e., Leymus chinensis and Phragmites communis, in a temperate meadow in northeastern China. Warming significantly increased the chlorophyll content and soluble sugars in L. chinensis but had no impact on the carotenoid and fructose contents. N addition caused a significant increase in the carotenoid and fructose contents. Warming and N addition had little impact on the photosynthetic products of P. communis. Warming caused significant decreases in the N and phosphorus (P) concentrations and significantly increased the carbon (C):P and N:P ratios of L. chinensis, but not the C concentration or the C:N ratio. N addition significantly increased the N concentration, C:P and N:P ratios, but significantly reduced the C:N ratio of L. chinensis. Warming significantly increased P. communis C and P concentrations, and the C:N and C:P ratios, whereas N addition increased the C, N and P concentrations but had no impact on the stoichiometric variables. This study suggests that both warming and N addition have direct impacts on plant photosynthates and elemental stoichiometry, which may play a vital role in plant-mediated biogeochemical cycling in temperate meadow ecosystems.     

Vertical Redistribution of Soil Organic Carbon Pools After Twenty Years of Nitrogen Addition in Two Temperate Coniferous Forests

Nitrogen (N) inputs from atmospheric deposition can increase soil organic carbon (SOC) storage in temperate and boreal forests, thereby mitigating the adverse effects of anthropogenic CO₂ emissions on global climate. However, direct evidence of N-induced SOC sequestration from low-dose, long-term N addition experiments (that is, addition of < 50 kg N ha⁻¹ y⁻¹ for > 10 years) is scarce worldwide and virtually absent for European temperate forests. Here, we examine how tree growth, fine roots, physicochemical soil properties as well as pools of SOC and soil total N responded to 20 years of regular, low-dose N addition in two European coniferous forests in Switzerland and Denmark. At the Swiss site, the addition of 22 kg N ha⁻¹ y⁻¹ (or 1.3 times throughfall deposition) stimulated tree growth, but decreased soil pH and exchangeable calcium. At the Danish site, the addition of 35 kg N ha⁻¹ y⁻¹ (1.5 times throughfall deposition) impaired tree growth, increased fine root biomass and led to an accumulation of N in several belowground pools. At both sites, elevated N inputs increased SOC pools in the moderately decomposed organic horizons, but decreased them in the mineral topsoil. Hence, long-term N addition led to a vertical redistribution of SOC pools, whereas overall SOC storage within 30 cm depth was unaffected. Our results imply that an N-induced shift of SOC from older, mineral-associated pools to younger, unprotected pools might foster the vulnerability of SOC in temperate coniferous forest soils.

     

Effects of Long-Term Nitrogen Addition and Atmospheric Nitrogen Deposition on Carbon Accumulation in Picea sitchensis Plantations

This study aimed to assess the combined effects of long-term nitrogen (N) supply and nitrogen deposition (N _dep) on carbon (C) accumulation within Sitka spruce [Picea sitchensis (Bong.) Carr.] plantations in Scotland. Six study sites established from 1970 to 1982 were periodically N-fertilized, monitored over time and commonly surveyed in 2010. Soil, aboveground biomass, and ground vegetation C stock changes were analyzed; aboveground C stocks were correlated with total additional N experienced at each site, that is, the sum of experimental N supply (N _add) and site-specific accumulated N _dep from 1900 to 2010. Results showed a positive N effect on aboveground tree C stock and no decline in tree growth was observed either during fertilization or after the latest N addition. The amount of C in litter was significantly higher in experimentally N-treated plots, whereas the amount of C in understory vegetation was higher in control plots. Pooling all the compartments (that is, understory vegetation, litter, soil, and tree biomass) the total ecosystem C content was estimated for each site, and at most sites a higher C stock was estimated for N-treated plots. Differences in aboveground C accumulation rates between treated and control plots were lower at sites with high levels of accumulated N _dep. Our results indicate that site-specific accumulated N _dep should be considered to understand tree growth responses to N fertilization.     

Nitrogen Fixation and Nitrogen Assimilation in a Temperate Saline Ecosystem

As nitrogen is known to be a limiting factor for plant growth, we were interested in the relationship between soil microbial activity and the nitrogen assimilation of 5 different halophytes from 4 saline sites near the lake “Neusiedlersee”, Austria. The following were studied between May and October 1985: nitrogen fixation (¹⁵N₂ and acetylene reduction): N-mineralization; several soil characteristics and in vivo nitrate reductase activity of roots and shoots of these plants. NO^?₃, org. N- and carboxylate contents of both roots and shoots, as well as the effect of NO^?₃-fertilization on the amounts of these substances, were determined on plants growing in the field during a 3-day period in September 1985. Fertilization led to a decrease in acetylene reduction activity at most sites, and an increase in the nitrate reductase activity of the shoots of all plants. Overall, carboxylate and organic nitrogen contents of these halophytes did not change in response to fertilization. Only in the roots of Aster tripolium and Atriplex hastata was there a marked increase in the nitrate reductase activity in response to fertilization. Species growing at the same site, such as Plantago maritima and Lepidium crassifolium showed contrasting levels of assimilatory activity. Apparent low rates of ammonification and nitrification were detected in soils from the 4 sites. The results are discussed in relation to the nitrogen and carbon economies of the microorganisms and plants.     

Long-Term Nitrogen Additions and Nitrogen Saturation in Two Temperate Forests

This article reports responses of two different forest ecosystems to 9 years (1988–96) of chronic nitrogen (N) additions at the Harvard Forest, Petersham, Massachusetts. Ammonium nitrate (NH₄NO₃) was applied to a pine plantation and a native deciduous broad-leaved (hardwood) forest in six equal monthly doses (May–September) at four rates: control (no fertilizer addition), low N (5 g N m^-2 y^-1), high N (15 g N m^-2 y^-1), and low N + sulfur (5 g N m^-2 y^-1 plus 7.4 g S m^-2 y^-1). Measurements were made of net N mineralization, net nitrification, N retention, wood production, foliar N content and litter production, soil C and N content, and concentrations of dissolved organic carbon (DOC) and nitrogen (DON) in soil water. In the pine stand, nitrate losses were measured after the first year of additions (1989) in the high N plot and increased again in 1995 and 1996. The hardwood stand showed no significant increases in nitrate leaching until 1995 (high N only), with further increases in 1996. Overall N retention efficiency (percentage of added N retained) over the 9-year period was 97–100% in the control and low N plots of both stands, 96% in the hardwood high N plot, and 85% in the pine high N plot. Storage in aboveground biomass, fine roots, and soil extractable pools accounted for only 16–32% of the added N retained in the amended plots, suggesting that the one major unmeasured pool, soil organic matter, contains the remaining 68–84%. Short-term redistribution of ¹⁵N tracer at natural abundance levels showed similar division between plant and soil pools. Direct measurements of changes in total soil C and N pools were inconclusive due to high variation in both stands. Woody biomass production increased in the hardwood high N plot but was significantly reduced in the pine high N plot, relative to controls. A drought-induced increase in foliar litterfall in the pine stand in 1995 is one possible factor leading to a measured increase in N mineralization, nitrification, and nitrate loss in the pine high N plot in 1996. Received 2 April 1999; Accepted 29 October 1999.     

模拟放牧斑块与氮素添加对半干旱草原群落植物生长的影响(英文)

放牧时,动物采食及其排泄物会影响植物的生长,但动物采食及其排泄物的空间异质性可能会影响这种效应。在位于我国北方典型农牧交错区的内蒙古多伦县,我们研究了模拟放牧斑块和施氮肥对植物生长的影响,实验采用模拟放牧采食斑块(刈割半径分别为0、10、20、40和80 cm)和土壤施氮(分别为0、5、10、20 g N/m2)两种处理,植物地上部收获后分为绿体和立枯两部分,并分析其含氮量。结果表明,刈割降低了植物的生物量(41.5％),而施氮可增加生物量(57.8％)。刈割对植物生长的抑制作用在面积最小又施肥的斑块上表现更明显。土壤施氮可以促进植物生长并且影响刈割效应。同时植物的绿-枯比随施氮水平的增加而增加,因此氮会延迟植物的衰老。以上结果表明,刈割(模拟动物采食)斑块的大小会影响草原植物的生长,土壤施氮(模拟动物尿氮)可以提高草原生态系统的初级生产力,并且影响刈割效应。     

The Inhibitory Effects of Nitrogen Deposition on Asymbiotic Nitrogen Fixation are Divergent Between a Tropical and a Temperate Forest

Ecosystems - Asymbiotic nitrogen (N) fixation (ANF) is an important source of N in pristine forests and is predicted to decrease with N deposition. Previous studies revealing N fixation in response...     

Effects of Water and Nitrogen Addition on Ecosystem Carbon Exchange in a Meadow Steppe

A changing precipitation regime and increasing nitrogen deposition are likely to have profound impacts on arid and semiarid ecosystem C cycling, which is often constrained by the timing and availability of water and nitrogen. However, little is known about the effects of altered precipitation and nitrogen addition on grassland ecosystem C exchange. We conducted a 3-year field experiment to assess the responses of vegetation composition, ecosystem productivity, and ecosystem C exchange to manipulative water and nitrogen addition in a meadow steppe. Nitrogen addition significantly stimulated aboveground biomass and net ecosystem CO₂ exchange (NEE), which suggests that nitrogen availability is a primary limiting factor for ecosystem C cycling in the meadow steppe. Water addition had no significant impacts on either ecosystem C exchange or plant biomass, but ecosystem C fluxes showed a strong correlation with early growing season precipitation, rather than whole growing season precipitation, across the 3 experimental years. After we incorporated water addition into the calculation of precipitation regimes, we found that monthly average ecosystem C fluxes correlated more strongly with precipitation frequency than with precipitation amount. These results highlight the importance of precipitation distribution in regulating ecosystem C cycling. Overall, ecosystem C fluxes in the studied ecosystem are highly sensitive to nitrogen deposition, but less sensitive to increased precipitation.     

Long-Term Nitrogen Addition Leads to Loss of Species Richness Due to Litter Accumulation and Soil Acidification in a Temperate Steppe

Background

Although community structure and species richness are known to respond to nitrogen fertilization dramatically, little is known about the mechanisms underlying specific species replacement and richness loss. In an experiment in semiarid temperate steppe of China, manipulative N addition with five treatments was conducted to evaluate the effect of N addition on the community structure and species richness.

Methodology/Principal Findings

Species richness and biomass of community in each plot were investigated in a randomly selected quadrat. Root element, available and total phosphorus (AP, TP) in rhizospheric soil, and soil moisture, pH, AP, TP and inorganic N in the soil were measured. The relationship between species richness and the measured factors was analyzed using bivariate correlations and stepwise multiple linear regressions. The two dominant species, a shrub Artemisia frigida and a grass Stipa krylovii, responded differently to N addition such that the former was gradually replaced by the latter. S. krylovii and A. frigida had highly-branched fibrous and un-branched tap root systems, respectively. S. krylovii had higher height than A. frigida in both control and N added plots. These differences may contribute to the observed species replacement. In addition, the analysis on root element and AP contents in rhizospheric soil suggests that different calcium acquisition strategies, and phosphorus and sodium responses of the two species may account for the replacement. Species richness was significantly reduced along the five N addition levels. Our results revealed a significant relationship between species richness and soil pH, litter amount, soil moisture, AP concentration and inorganic N concentration.

Conclusions/Significance

Our results indicate that litter accumulation and soil acidification accounted for 52.3% and 43.3% of the variation in species richness, respectively. These findings would advance our knowledge on the changes in species richness in semiarid temperate steppe of northern China under N deposition scenario.     

Correction to: Vertical Redistribution of Soil Organic Carbon Pools After Twenty Years of Nitrogen Addition in Two Temperate Coniferous Forests

Ecosystems - In the article by Forstner et al. (2018), the surnames of co-authors Katharina M. Keiblinger and Patrick Schleppi were misspelled. We apologize and ask readers to cite the corrected...