Tree growth and soil acidification in response to 30 years of experimental nitrogen loading on boreal forest

Relations among nitrogen load, soil acidification and forest growth have been evaluated based on short‐term (<15 years) experiments, or on surveys across gradients of N deposition that may also include variations in edaphic conditions and other pollutants, which confound the interpretation of effects of N per se. We report effects on trees and soils in a uniquely long‐term (30 years) experiment with annual N loading on an un‐polluted boreal forest. Ammonium nitrate was added to replicated (N=3) 0.09 ha plots at two doses, N1 and N2, 34 and 68 kg N ha^?1 yr^?1, respectively. A third treatment, N3, 108 kg N ha^?1 yr^?1, was terminated after 20 years, allowing assessment of recovery during 10 years. Tree growth initially responded positively to all N treatments, but the longer term response was highly rate dependent with no gain in N3, a gain of 50 m³ ha^?1 stemwood in N2 and a gain of 100 m³ ha^?1 stemwood in excess of the control (N0) in N1. High N treatments caused losses of up to 70% of exchangeable base cations (Ca²⁺, Mg²⁺, K⁺) in the mineral soil, along with decreases in pH and increases in exchangeable Al³⁺. In contrast, the organic mor‐layer (forest floor) in the N‐treated plots had similar amounts per hectare of exchangeable base cations as in the N0 treatment. Magnesium was even higher in the mor of N‐treated plots, providing evidence of up‐lift by the trees from the mineral soil. Tree growth did not correlate with the soil Ca/Al ratio (a suggested predictor of effects of soil acidity on tree growth). A boron deficiency occurred on N‐treated plots, but was corrected at an early stage. Extractable NH₄⁺ and NO₃^?were high in mor and mineral soils of on‐going N treatments, while NH₄+ was elevated in the mor only in N3 plots. Ten years after termination of N addition in the N3 treatment, the pH had increased significantly in the mineral soil; there were also tendencies of higher soil base status and concentrations of base cations in the foliage. Our data suggest the recovery of soil chemical properties, notably pH, may be quicker after removal of the N‐load than predicted. Our long‐term experiment demonstrated the fundamental importance of the rate of N application relative to the total amount of N applied, in particular with regard to tree growth and C sequestration. Hence, experiments adding high doses of N over short periods do not mimic the long‐term effects of N deposition at lower rates.     

Mineralization of amino acids and its signs in nitrogen cycling of forest soil

Amino acid mineralization and its fate in soil have effects on soil nitrogen cycling. Here we used ¹⁵N-labeled alanine and methionine to study differences in their mineralization from soil organic nitrogen under 60% WHC (water holding capacity) and 90% WHC soil conditions. We found that the maximum mineralization rates were at the 24th hours for alanine and at the 5th hours for methionine, and about two times greater rates at 60% WHC than at 90% WHC. The half-live was 24–72 h for alanine and > 72 h for methionine. Half-lives of amino acids occurred sooner under 90% WHC than under 60% WHC. The results suggested that some kind of amino acids do lead the nitrogen cycling in a specific ecosystem or as a sign to trigger soil nitrogen cycling when land utilization was altered or disturbed severely by humans.     

氮沉降对森林植物的影响

综述了氮沉降对森林植物的影响。氮沉降对森林植物的影响主要表现在以下6个方面：(1)在一定量范围内的氮沉降有利于植物的光合作用,但过量后则会引起植物的光合速率下降;(2)当植物生长受氮限制时,在一定程度上的氮沉降增加植物生产力,但当氮过量后,氮沉降则使植物的生产力下降;(3)过量的氮沉降导致植物体各种营养元素含量的比例失衡;(4)氮沉降会改变植物的形态结构,集中表现为根／冠比减小;(5)氮沉降会增加植物对天然胁迫如干旱、病虫害和风的敏感性,减少其抵御能力;(6)氮沉降会改变植物组成和降低森林植物的多样性。     

Contribution of amino compounds to dissolved organic nitrogen in forest soils

Dissolved organic nitrogen (DON) may play an important role in plantnutrition and nitrogen fluxes in forest ecosystems. In spite of the apparentimportance of DON, there is a paucity of information concerning its chemicalcomposition. However, it is exactly this chemical characterization that isrequired to understand the importance of DON in ecosystem processes. Theprimaryobjective of this study was to characterize the distribution of free aminoacidsand hydrolyzable peptides/proteins in the DON fraction of Oa horizon leachatesalong an extreme edaphic gradient in northern California. Insitu soil solutions were extracted by centrifugation from Oahorizonscollected beneath Pinus muricata (Bishop pine) andCupressus pygmaea (pygmy cypress) on slightlyacidic/fertile and highly acidic/infertile sites. DON accounted for 77 to99% of the total dissolved nitrogen in Oa horizon leachates. Nitrogen infree amino acids and alkyl amines ranged from 0.04–0.07 mgN/L on the low fertility site to 0.45–0.49 mg N/L onthe high fertility site, and accounted for 1.5 to 10.6% of the DON fraction.Serine, glutamic acid, leucine, ornithine, alanine, aspartic acid andmethylamine were generally the most abundant free amino compounds. Combinedamino acids released by acid hydrolysis accounted for 48 to 74% of theDON, suggesting that proteins and peptides were the main contributor to DON inOa horizon leachates. Together, nitrogen from free andcombined amino compounds accounted for 59 to 78% of the DON. Most of theDON was found in the hydrophobic fraction, which suggests the presence ofprotein/peptide-polyphenol complexes or amino compounds associated withhumic substances. Because free and combined amino acids can be an importantnitrogen source for some plants, soil DON may play an important role in plantnutrition and ecosystem function.     

Nonlinear responses of ecosystem carbon fluxes to nitrogen deposition in an old-growth boreal forest

Nitrogen (N) deposition is known to increase carbon (C) sequestration in N-limited boreal forests. However, the long-term effects of N deposition on ecosystem carbon fluxes have been rarely investigated in old-growth boreal forests. Here we show that decade-long experimental N additions significantly stimulated net primary production (NPP) but the effect decreased with increasing N loads. The effect on soil heterotrophic respiration (Rh) shifted from a stimulation at low-level N additions to an inhibition at higher levels of N additions. Consequently, low-level N additions resulted in a neutral effect on net ecosystem productivity (NEP), due to a comparable stimulating effect on NPP and Rh, while NEP was increased by high-level N additions. Moreover, we found nonlinear temporal responses of NPP, Rh and NEP to low-level N additions. Our findings imply that actual N deposition in boreal forests likely exerts a minor contribution to their soil C storage.     

森林生态系统碳循环对全球氮沉降的响应

森林土壤和植被储存着全球陆地生态系统大约46%的碳,在全球碳平衡中起着非常重要的作用。过去几十年来,森林生态系统的碳循环和碳吸存受到了全球氮沉降的深刻影响,因为氮沉降改变了陆地生态系统的生产力和生物量积累。以欧洲和北美温带森林区域开展的研究为基础,综述了氮沉降对植物光合作用、土壤呼吸、土壤DOM及林木生长的影响特征和机理,探讨了森林生态系统碳动态对氮沉降响应的不确定性因素。热带森林C、N循环与大部分温带森林不同,人为输入的氮对热带生态系统过程的影响也可能不同,因此指出了在热带地区开展碳氮循环耦合研究的必要性和紧迫性。     

Incorporation of nitrate nitrogen into amino acids during the anaerobic germination of rice

Summary Incorporation of¹⁵NO₃-into amino acids was studied during the anaerobic germination of rice seeds. In treated coleoptiles, the label was incorporated into glutamine, glutamate, alanine,-aminobutyric acid (Gaba), arginine, aspartate and methionine. These findings are consistent with a primary incorporation of nitrate nitrogen into glutamine, glutamate and aspartate, and their further conversion to alanine, Gaba, arginine and methionine.     

Mass loss and nitrogen dynamics in decomposing acid forest litter in the Netherlands at increased nitrogen deposition

Litterbag experiments were carried out in five forest ecosystems in the Netherlands to study weight loss and nitrogen dynamics during the first two years of decomposition of leaf and needle litter. All forests were characterized by a relatively high atmospheric nitrogen input by throughfall, ranging from 22–55 kg N ha^–1 yr^–1.Correlation analysis of all seven leaf and needle litters revealed no significant relation between the measured litter quality indices (nitrogen and lignin concentration, lignin-to-nitrogen ratio) and the decomposition rate. A significant linear relation was found between initial lignin-to-nitrogen ratio and critical nitrogen concentration, suggesting an effect of litter quality on nitrogen dynamics.Comparison of the decomposition of oak leaves in a nitrogen-limited and a nitrogen-saturated forest suggested an increased nitrogen availability. The differences in capacities to retain atmospheric nitrogen inputs between these two sites could be explained by differences in net nitrogen immobilization in first year decomposing oak leaves: in the nitrogen-limited oak forest a major part (55%) of the nitrogen input by throughfall was immobilized in the first year oak leaf litter.The three coniferous forests consisted of two monocultures of Douglas fir and a mixed stand of Douglas fir and Scots pine. Despite comparable litter quality in the Douglas fir needles in all sites, completely different nitrogen dynamics were found.     

Estimation of nitrogen saturation on the basis of long-term fertilization experiments

The obvious changes in some properties of forest soil due to atmospheric nitrogen deposition under Finnish conditions were estimated on the basis of long-term fertilization experiments. The experiments were established during the years 1959–1965. Nitrogen fertilization was repeated three to four times. During the study period averaging 23 years, the cumulative amount of fertilizer nitrogen totalled about 400 kg N ha^-1. The main results are as follows. Nitrogen addition increased the quantity of organic matter in the humus layer, but has not clear effect on its quality. Nitrogen fertilization did not increase soil acidity.     

Remarks on the effects of nitrogen deposition to forest ecosystems

The effects of increased deposition of nitrogen compounds on forest sites are discussed based on literature data and own results from both earlier fertilization experiments and the ARINUS study area in the Black Forest. The influence on mycorrhiza is stated suppressive as well as stimulating so that no general conclusion can be drawn. The nitrogen nutrition status of coniferous forests is still sub-optimal over wide areas with a yearly deposition of 10–20 kg N ha^-1. Under considerable higher input rates the insufficient supply of other nutrients and imbalances in the nutritional status of trees are possible. When discussing nitrogen saturation of ecosystems, the nitrogen storage capacity of soils has to be considered as a decisive factor which varies from site to site. Any actual input/output balance is strongly influenced by the internal turnover processes resulting from former land use.     

Microbial activity and soil respiration under nitrogen addition in Alaskan boreal forest

Climate warming could increase rates of soil organic matter turnover and nutrient mineralization, particularly in northern high‐latitude ecosystems. However, the effects of increasing nutrient availability on microbial processes in these ecosystems are poorly understood. To determine how soil microbes respond to nutrient enrichment, we measured microbial biomass, extracellular enzyme activities, soil respiration, and the community composition of active fungi in nitrogen (N) fertilized soils of a boreal forest in central Alaska. We predicted that N addition would suppress fungal activity relative to bacteria, but stimulate carbon (C)‐degrading enzyme activities and soil respiration. Instead, we found no evidence for a suppression of fungal activity, although fungal sporocarp production declined significantly, and the relative abundance of two fungal taxa changed dramatically with N fertilization. Microbial biomass as measured by chloroform fumigation did not respond to fertilization, nor did the ratio of fungi : bacteria as measured by quantitative polymerase chain reaction. However, microbial biomass C : N ratios narrowed significantly from 16.0 ± 1.4 to 5.2 ± 0.3 with fertilization. N fertilization significantly increased the activity of a cellulose‐degrading enzyme and suppressed the activities of protein‐ and chitin‐degrading enzymes but had no effect on soil respiration rates or ¹⁴C signatures. These results indicate that N fertilization alters microbial community composition and allocation to extracellular enzyme production without affecting soil respiration. Thus, our results do not provide evidence for strong microbial feedbacks to the boreal C cycle under climate warming or N addition. However, organic N cycling may decline due to a reduction in the activity of enzymes that target nitrogenous compounds.     

Mycorrhizal status of herb-layer plants in a fertilized oak-pine forest

In the years 1986–1988 investigations were carried out on the influence of fertilizers on the mycorrhizal status of herb-layer plants of a mixed oak-pine forest in the Niepolomice Forest (Southern Poland). The site enrichment with N, P, K altered the frequency of mycorrhizal infection in most species investigated and decreased the percentage of colonized root cells. Some species disappear after fertilization (e.g. Vaccinium myrtillus L.). Among the species vigorously expanding after the treatment, mainly nonmycorrhizal plants (e.g. Festuca gigantea (L.) Vill.) were found. Some species (e.g. Milium effusum L.) favourably reacting to mineral fertilization, lost their mycorrhizae.     

川南天然常绿阔叶林土壤酶活性特征及其对模拟N沉降的响应

通过原位进行低氮(LN,50 kg N·hm-2·a-1)、中氮(MN,100 kg N·hm-2·a-1)和高氮(HN,150 kg N·hm-2·a-1)处理,研究了川南天然常绿阔叶林土壤酶活性特征及其对模拟N沉降的响应.结果表明,森林土壤过氧化氢酶、脲酶、纤维素酶和蔗糖酶的活性在垂直分布上均表现为0～10 cm土层高于10～20 cm土层;各种土壤酶活性的季节变化明显,但土壤酶活性还因取样时间和土层不同而异.其中,0～10 cm和10～20 cm土层中过氧化氢酶活性高峰均出现在秋季(92 d);0～10 cm土层中脲酶活性高峰出现在秋季,而10～20 cm土层中脲酶活性高峰出现在冬季(183 d);0～10 cm和10～20 cm土层中土壤纤维素酶活性高峰则出现在翌年春季(274 d);0～10 cm土层中蔗糖酶活性有两个高峰,分别出现在秋季和翌年春季;而10～20 cm土层中蔗糖酶活性只有一个高峰,出现在翌年春季.总体上,常绿阔叶林中不同土壤酶活性对N沉降的响应有所差异.其中,N沉降降低了常绿阔叶林土壤纤维素酶和过氧化氢酶活性,增加了土壤脲酶和蔗糖酶活性.不同浓度N处理间酶活性表现出了不同程度的差异性,但其变化的规律还不明显.     

Nitrogen transformations in two nitrogen saturated forest ecosystems subjected to an experimental decrease in nitrogen deposition

Nitrogen transformations were studied in the forest floor and mineral soil (0–5 cm) of a Douglas fir forest (Pseudotsuga menziesii (Mirb.) Franco.) and a Scots pine forest (Pinus sylvestris L.) in the Netherlands. Curren nitrogen depositions (40 and 56 kg N ha^-1 yr^-1, respectively) were reduced to natural background levels (1–2 kg N ha^-1 yr^-1) by a roof construction. The study concentrated on rates and dynamic properties of nitrogen transformations and their link with the leaching pattern and nitrogen uptake of the vegetation under high and reduced nitrogen deposition levels. Results of an in situ field incubation experiment and laboratory incubations were compared. No effect of the reduced N deposition on nitrogen transformations was found in the Douglas fir forest. In the Scots pine forest, however, during some periods of the year nitrogen transformations were significantly decreased under the low nitrogen deposition level. At low nitrogen inputs a net immobilization occurred during most of the year leading to a very small net mineralization for the whole year. In laboratory and in individual field plots nitrogen transformations were negatively correlated with initial inorganic nitrogen concentrations. Nitrogen budget estimates showed that nitrogen transformations were probably underestimated by the in situ incubation technique. Nevertheless less nitrogen was available for plant uptake and leaching at the low deposition plots.     

模拟氮沉降对温带阔叶红松林地土壤氮素净矿化的影响

以长白山阔叶红松混交林为研究对象,于2006—2008年原位模拟不同形态氮((NH4)2SO4、NH4Cl和KNO3)沉降水平(22.5和45kgN·hm-2·a-1),利用树脂芯法技术(resin-core incubation technique)测定了表层(有机层0～7cm)和土层(0～15cm)土壤氮素净矿化、净氨化和净硝化通量的季节和年际变化规律。同时,结合前人报道的有关林地碳、氮过程及其环境变化影响的结果,力求有效预估森林生态系统中氮素年矿化通量对大气氮沉降量和水热条件等因子变化的响应。结果表明,长白山阔叶红松林地土壤氮素年净矿化通量为1.2～19.8kgN·hm-2·a-1,2008年不同深度的土壤氮素年净矿化通量均显著高于2006和2007年(P<0.05)。随着模拟氮沉降量增加,土壤氮素净矿化通量也随之增加,尤其外源NH4+-N输入对净矿化通量的促进作用更为明显(P<0.05),但随着施肥年限的延长,这种促进作用逐渐减弱。与林地0～15cm土壤相比,氮沉降增加对0～7cm有机层氮素净氨化和净矿化通量的促进作用更为明显,尤其NH4Cl处理的促进作用更大。结合前人报道的野外原位观测结果,土壤氮素年净矿化通量随氮素沉降量的增加而增大,氮沉降量对不同区域森林土壤氮素净矿化通量的贡献率约为52%;氮沉降量(x1)和pH值(x2)可以解释区域森林土壤氮素年净矿化通量(y)变化的70%(y=0.54x1-18.38x2-109.55,R2=0.70,P<0.0001)。前人研究结果仅提供区域年均温度,未考虑积温的影响,这可能是造成年净矿化通量与温度无关的原因。今后的研究工作应该加强区域森林土壤积温观测,进而更加准确地预估森林土壤氮素的年净矿化通量。     

An attempt to predict long-term effects of atmospheric nitrogen deposition on the yield of Norway spruce stands (Picea abies (L.) Karst.) in southwestern Sweden

Long-term field experiments in Norway spruce stands on fertile sites (site indices 27–35 m) in southwestern Sweden were analysed with respect to volume increment. Three treatments were included (0=No fertilization, N = Fertilization with N, NP = Fertilization with N and P).Volume growth was monitored for 18 years in 10 blocks. No significant differences in annual volume increment between the treatments were detected. Volume increments in the N treatment were 97%, 99% and 107% as high as those in the 0 treatment for the periods 1–5, 6–10 and 11–15 years after the first fertilization. Corresponding values for the NP treatment were 104%, 108% and 110%, indicating that P has a small positive effect.The amount of N-fertilization would correspond to an annual N deposition of 20 kg ha^-1 during the next 30 years in southwestern Sweden. For this period, the results imply that this N deposition would not affect the growth of Norway spruce stands on fertile sites.     

Mineral nitrogen and microbial dynamics in the forest floor of clearcut or partially harvested successional boreal forest stands

The effects of clearcut and partial harvesting of early-seral trembling aspen plots were compared to conventional clearcut harvesting in mid-seral mixedwood and late-seral conifer plots. Twice a year, for three consecutive years, we assessed mineral N and microbial dynamics in the forest floor of these plots to test three hypotheses related to the higher litter quality of aspen leaves and to the sustained inputs of available C on partially harvested plots: (1) the post-clearcutting mineral N flush and the net [(NO₃^–): (NO₃^– + NH₄⁺)] production ratio (RNI) are higher in aspen plots than in black spruce plots, with intermediate values occurring in mixedwood plots; (2) net N mineralization rates in aspen plots are higher in spring than in autumn; and (3) compared to clearcutting, partial harvesting reduces potential ammonification and nitrification rates. Initial NH₄⁺ and NO₃^– concentrations respectively ranged between 1.7–4.4 and 0.2–1.5 g N kg^–1 N_total, net ammonification and nitrification rates (30 d incubations) respectively ranged between 5.3–17.8 and 0.1–27.6 g N kg^–1 N_total, basal respiration ranged between 20.9–38.9 mg CO₂-C kg^–1 h^–1, and microbial biomass ranged between 6.1–8.7 g C_mic kg^–1. Although clearcutting increased NO₃^– concentrations in aspen plots, the balance of our results did not support our first hypothesis, because NH₄⁺ concentrations increased in conifer plots only, potential ammonification was unaffected by clearcutting, potential nitrification increased in mixedwood plots only, and RNI increased in all plots. In each seral stage, basal respiration, microbial biomass, and metabolic quotient either increased or were unaffected by clearcutting, suggesting that increases in RNI after disturbance were not related to lower microbial immobilisation of NO₃^– due to lower available C. Forest floors in mid-seral mixedwood plots exhibited a distinct combination of mineral N and microbial properties, suggesting that the functional richness of the forest is enhanced not only by the number of species, but also by the diversity of assemblages that are present. Results supported our second hypothesis and showed, furthermore, that net N mineralization in conifer stands is greater in autumn than in spring. Partial harvesting in aspen stands resulted in lower potential mineralization of N and lower RNI, compared to clearcutting. Further lysimetry studies are needed to confirm whether partial harvesting mitigates NO₃^– leaching following disturbance.