Effects of simulated N deposition on understorey vegetation of a boreal coniferous forest

1. We report the short-term effects of simulated nitrogen (N) deposition on the understorey of a boreal forest in northern Sweden. Doses of ¹⁵N double-labelled NH₄NO₃ (0·5, 12·5, 25·0 and 50·0 kg N ha^–1) were applied to 1 m² plots in the summer of 1995 and plants were harvested the following autumn.
2. No significant treatment effects were found in either above- or below-ground biomass which was distributed as follows: the ericaceous shrub Vaccinium myrtillus contributed 76%, the grass Deschampsia flexuosa 4%, and the bryophytes Dicranum majus and Pleurozium schreberi 20%.
3. The recovery of applied N in these species was 24, 27, 27 and 32% of the 0·5, 12·5, 25·0 and 50·0 kg N ha^–1 applications, respectively, and thus the recovery increased with the N dose.
4. In the 0·5 kg N ha^–1 treated plots, the highest concentrations of fertilizer-derived ¹⁵N were found in the bryophytes, while in plots given the three higher N applications, leaves of D . flexuosa had the highest concentrations.
5. N application resulted in elevated concentrations of free amino acid N, which indicate increased storage of N in plant tissues. Furthermore, the N application resulted in increased damage to V . myrtillus by natural enemies. The severity of disease caused by two foliar parasitic fungi showed a clear numerical response to N treatment, as did the amount of herbivory as a result of Lepidoptera larvae.     

Moose density and habitat productivity affects reproduction,growth and species composition in field layer vegetation

Question: What is the effect of a gradient in moose density on reproduction, growth and functional composition of the field layer vegetation in a boreal forest, and how is this effect modified by habitat productivity? Location: Northwest of Umeå, Västerbotten, northern Sweden. Methods: Field layer vegetation was surveyed in an experimental setup with simulation of three different moose densities and a control in eight study sites along a gradient of habitat productivity. Results: We found that increased moose density led to decreased cover and reproductive effort of a browsed dwarf shrub (bilberry, Vaccinium myrtillus L.) and increased cover and reproductive effort of a non‐browsed graminoid (wavy hair‐grass, Avenella flexuosa (L.) Drejer). Increased moose density led to increased light availability and probably reduced competition from V. myrtillus. Total reproductive effort in the field layer vegetation increased, height decreased and cover of light‐demanding species and graminoids increased with increasing moose density. The effects of moose density were modified by the productivity gradient, leading to a higher relative increase in light availability and reproductive effort in highly productive areas than in low productive areas. Conclusions: Increased light availability was an important indirect effect of moose density, leading to less competition for light and a shift towards early successional species. The effect of moose density on light availability was modified by habitat productivity, leading to stronger relative effects in highly productive areas than in low productive areas.     

Individual and combined effects of disturbance and N addition on understorey vegetation in a subarctic mountain birch forest

Questions: What are the effects of repeated disturbance and N‐fertilization on plant community structure in a mountain birch forest? What is the role of enhanced nutrient availability in recovery of understorey vegetation after repeated disturbance? How are responses of soil micro‐organisms to disturbance and N‐fertilization reflected in nutrient allocation patterns and recovery of understorey vegetation after disturbance? Location: Subarctic mountain birch forest, Finland. Methods: We conducted a fully factorial experiment with annual treatments of disturbance (two levels) and N‐fertilization (four levels) during 1998–2002. We monitored treatment effects on above‐ground plant biomass, plant community structure and plant and soil nutrient concentrations. Results: Both disturbance and N‐fertilization increased the relative biomass of graminoids. The increase of relative biomass of graminoids in the disturbance treatment was over twice that of the highest N‐fertilization level, and N‐fertilization further increased their relative biomass after disturbance. As repeated disturbance broke the dominance of evergreen dwarf shrubs, it resulted in a situation where deciduous species, graminoids and herbs dominated the plant community. Although relative biomass of deciduous dwarf shrubs declined with N‐fertilization, it did not cause a shift in plant community structure, as evergreen dwarf shrubs remained dominant. Both disturbance and N‐fertilization increased the N concentration in vascular plants, whereas microbial biomass N and C were not affected by the treatments. Concentrations of NH₄⁺, dissolved organic N (DON) and dissolved organic C (DOC) increased in the soil after N‐fertilization, whereas concentrations of NH₄⁺ and DON decreased after disturbance. Conclusions: Disturbances caused by e.g. humans or herbivores contribute more to changes in the understorey vegetation structure than increased levels of N in subarctic vegetation. Fertilization accelerated the recovery potential after repeated disturbance in graminoids. Microbial activities did not limit plant growth.     

Regeneration strategies of dominant boreal forest dwarf shrubs in response to selective removal of understorey layers

Abstract. The deciduous Vaccinium myrtillus and the evergreen Vaccinium vitis‐idaea were subjected to five removal treatments of understorey layers: control, removal of the moss layer, removal of the field layer, removal of both moss and field layers and removal of moss, field and humus layers. A second factor, sowing, was included to investigate sexual reproduction after disturbance. Density of new ramets and seedlings and growth of annual shoots were studied for the first two growing seasons, whereas cover was measured for five growing seasons after disturbance treatment. Initially, vegetative production of new ramets and species cover increased rapidly in all disturbed plots, except for the most severe treatment, in which production of new ramets was virtually absent throughout the study. Full recovery following removal of the field layer only or both field and moss layers was reached after four years for V. myrtillus. V. vitis‐idaea recovered more quickly, after one year (removal of field layer only) and four years (removal of field and moss layers). The relative growth of V. myrtillus and V. vitis‐idaea increased in the latter treatment in terms of production of annual shoots and length of annual shoots, respectively. Seedling density increased after sowing in the most severe treatment. The results underscore the importance of vegetative growth for recovery of these species at moderate‐level disturbances. The high rate of sexual reproduction in the most severe treatment implies that strong mechanical disturbance is needed to enhance the establishment of new genotypes in these species.     

Photosynthesis of ground vegetation in different aged pine forests: Effect of environmental factors predicted with a process‐based model

Question: How do stand age and environmental factors affect the species‐specific photosynthesis of ground vegetation? Location: Five different aged pine forests in Southern Finland. Methods: We measured photosynthesis of common species of ground vegetation during the growing season of 2006. Results: The measured vascular species, especially those with annual leaves, had a clear seasonal cycle in their measured photosynthetic activity (P_maxⁱ). A simple model that uses site‐specific temperature history, soil moisture and recent frost as input data was able to predict the changes in photosynthetic activity in dwarf shrubs with perennial leaves. The P_maxⁱ values of mosses did not have a clear seasonal cycle, but low values occurred after rain‐free periods and high values after precipitation. We modified the model for mosses and included temporary rain events. The model was able to predict most of the large changes in P_maxⁱ of mosses resulting from varying weather events but there was still some uncertainty, which was probably due to difficulties in measuring fluxes over a moss population. Conclusions: Temperature history, recent frosts and soil moisture determine the changes in P_maxⁱ of dwarf shrubs with perennial leaves. The P_maxⁱ of mosses depends mostly on recent precipitation.     

Slow response of soil organic matter to the reduction in atmospheric nitrogen deposition in a Norway spruce forest

Global nitrogen (N) deposition rates in terrestrial environments have quadrupled since preindustrial times, causing structural and functional changes of ecosystems. Different emission reduction policies were therefore devised. The aim of our study was to investigate if, and over what timescale, processes of soil organic matter (OM) transformation respond to a decline in atmospheric N deposition. A N‐saturated spruce forest (current N deposition: 34 kg ha^?1 yr^?1; critical N load: 14 kg ha^?1 yr^?1), where N deposition has been reduced to 11.5 kg ha^?1 yr^?1 since 1991, was studied. Besides organic C and organic and inorganic N, noncellulosic carbohydrates, amino sugars and amino acids were determined. A decline in organic N in litter indicated initial effects at plant level. However, there were no changes in biomarkers upon the reduction in N deposition. In addition, inorganic N was not affected by reduced N deposition. The results showed that OM cycling and transformation processes have not responded so far. It was concluded that no direct N deposition effects have occurred due to the large amount of stored organic N, which seems to compensate for the reduction in deposited N. Obviously, the time span of atmospheric N reduction (about 14.5 years) is too short compared with the mean turnover time of litter to cause indirect effects on the composition of organic C and N compounds. It is assumed that ecological processes, such as microbial decomposition or recycling of organic N and C, react slowly, but may start within the next decade with the incorporation of the new litter.     

1954—2005年中国北方针叶林分布区的气候变化特征

基于中国北方针叶林(兴安落叶松林)分布区8个气象观测站的气象资料,分析了1954—2005年气温和降水的变化特征.结果表明：研究期间,中国北方针叶林分布区的气温以0.38 ℃·(10 a)^-1的速度上升,远大于全球近50年来0.13 ℃·(10 a)^-1的平均增温速率.尽管夏、秋季的气温呈上升趋势,但不显著;而冬、春季的增温显著(P<0.01);最高年平均气温(0.37 ℃·(10 a)^-1)与最低年平均气温(0.54 ℃·(10 a)^-1)均呈极显著的增加趋势(P<0.01).降水量年际间波动较大,但没有明显的变化趋势;各季节降水量也没有明显的变化规律,其中春、秋、冬季的降水日数有增加趋势,但没有达到显著水平,而夏季的降水日数呈显著减少趋势(P<0.05);各季降水强度均呈增加趋势,其中夏季(P<0.05)和冬季(P<0.01)的变化达到了显著水平.     

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.     

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.     

Changes in the composition of rainwater upon passage through the canopies of trees and of ground vegetation in a Dutch oak-birch forest

The composition of canopy throughfall water in an oak-birch woodland, heavily affected by atmospheric deposition of N and S, changed markedly upon contact with the above-ground parts of the ground vegetation, which consisted mainly of bracken. The fluxes of nitrate and H⁺ decreased, simultaneously with an increase in the flux of bicarbonate, indicative of above-ground uptake of nitrate by the ground vegetation. This above-ground assimilation takes place in spite of abundant availability of inorganic nitrogen in the root zone of the ground vegetation. Fluxes of phosphate were somewhat lower, and those of ammonium somewhat higher in throughfall of the ground vegetation than in that of the tree layer. Although those differences were not statistically significant, they do suggest assimilation of some P and extra net dry deposition of atmospheric ammonia below the tree canopies.     

Seasonal Changes of Nitrogen Storage Compounds in a Rhizomatous Grass Calamagrostis epigeios

The seasonal dynamics in content and distribution of N-rich compounds between overwintering organs of Calamagrostis epigeios were examined. Samples were taken both from plants grown in natural conditions and in containers with controlled nutrient supply. There were significant changes in content of nitrate, free amino acids and soluble protein in all investigated plant parts during the course of a year. Amino acids showed both the highest maximum and seasonal fluctuation among the all N compounds observed and, therefore, appear to have a central role in N storage. Their content rises in the autumn, remains stable during winter and declines quickly at the beginning of spring. The most abundant amino acids in the end of winter storage period - asparagine, arginine and glutamine - constituted about 90 % of N in fraction of free amino acids. The portion of N stored in soluble proteins, however, was considerably smaller compare to both amino acids and nitrate. The amount of N stored in rhizomes of C. epigeios was smaller than in roots and stubble base before the onset of spring re-growth. This indicates that roots and stubble base are particularly important for winter N storage in this species.     

森林土壤酶对环境变化的响应研究进展

全球气候变化已是不争的事实,对陆地生态系统特别是森林生态系统物质循环将产生显著的影响。土壤酶是森林土壤物质循环的主要限制因素之一,对气候变化的响应近年来受到广泛关注。由于森林土壤酶对全球气候变化的响应研究是预测未来环境变化对森林生态系统过程影响的关键,因此,着重综述了森林土壤酶对环境变化尤其是全球变暖和氮沉降响应方面的研究,并分析了未来研究的主要方向。环境变化会引起土壤p H、水分及其营养成分的变化,而这些变化会反作用于土壤酶的活性和稳定性。森林土壤酶对增温的响应,不仅与酶的种类以及增温的温度范围和持续时间有关,还与土壤类型有关,是多种因子综合作用的结果。森林土壤酶对氮添加的响应与林分类型和土层类型有关,受复合氮的影响更大。建议未来的研究应加强酶的基本性质对环境变化的响应研究,注重林分类型、土层类型导致的差异,强化多因素的交互作用,并进行长期、综合的观测。     

模拟氮沉降对华西雨屏区天然常绿阔叶林土壤微生物生物量碳和氮的影响

为理解模拟氮沉降对华西雨屏区天然常绿阔叶林土壤微生物生物量碳(MBC)和氮(MBN)的影响,通过一年野外模拟氮(NH₄NO₃)沉降试验,氮沉降水平分别为对照(CK, 0 g N·m^-2·a^-1)、低氮沉降(L, 5 g N·m^-2·a^-1)、中氮沉降(M, 15 g N·m^-2·a^-1)和高氮沉降(H, 30 g N·m^-2·a^-1),研究了氮沉降对天然常绿阔叶林土壤MBC和MBN的影响.结果表明: 氮沉降显著降低了0～10 cm土层MBC和MBN,且随氮沉降量的增加,下降幅度增大;L和M处理对10～20 cm土层MBC和MBN无显著影响,H处理显著降低了10～20 cm土层土壤MBC和MBN;氮沉降对MBC和MBN的影响随土壤深度的增加而减弱.MBC和MBN具有明显的季节变化,在0～10和10～20 cm土层均表现为秋季最高,夏季最低.0～10和10～20 cm土层土壤微生物生物量C/N分别介于10.58～11.19和9.62～12.20,表明在华西雨屏区天然常绿阔叶林土壤微生物群落中真菌占据优势.