Inferred effects of cloud deposition on forest floor nutrient cycling and microbial properties along a short elevation gradient

Cloud water deposition often increases with elevation, and it is widely accepted that this cloud water increases acid loading to upland forest ecosystems. A study was undertaken in south-eastern Quebec to determine if a 250 m elevation gradient (i.e. 420-665 m), along a uniform sugar-maple stand on the slope of Mount Orford, corresponded to a pH gradient in the forest floor and to predictable changes in soil nutrient availability and microbial properties. Precipitation data from a nearby study, and a photographic survey, provided presumptive evidence that this elevation gradient corresponded to a strong gradient in cloud water deposition. Forest floor temperature did not differ significantly across elevations. Forest floor moisture content was significantly higher, whereas pH and exchangeable Ca and Mg were significantly lower, at the higher elevations. Average seasonal net nitrification rates, determined by long-term laboratory incubations, did not differ significantly across elevations, whereas average seasonal net ammonification rates were significantly higher at higher elevations. Basal respiration rates and microbial biomass did not differ significantly across elevations, but metabolic quotient was significantly higher at higher elevations indicating possible environmental stress on forest floor microbial communities due to cloud water deposition. Anaerobic N mineralisation rates were significantly higher at higher elevations suggesting that N-limited microbial communities frequently exposed to cloud cover can be important short-term sinks for atmospheric N, thereby contributing to increase the active-N fraction of forest floors. We conclude that, where no significant changes in vegetation or temperature occur, elevation gradients can still be used to understand the spatial variability of nutrient cycles and microbial properties.     

Gaseous nitrogen losses from a forest site in the North Tyrolean Limestone Alps

Microorganisms are responsible for the mineralisation of organic nitrogen in soils. NH ⁺₄ can be further oxidised to NO₃ ⁻ during nitrification and NO₃ ⁻ can be reduced to gaseous nitrogen compounds during denitrification. During both processes, nitrous oxide (N₂O), which is known as greenhouse gas, can be lost from the ecosystem.

The aim of this study was to quantify N₂O emissions and the internal microbial N cycle including net N mineralisation and net nitrification in a montane forest ecosystem in the North Tyrolean Limestone Alps during an 18-month measurement period and to estimate the importance of these fluxes in comparison with other components of the N cycle. Gas samples were taken every 2 weeks using the closed chamber method. Additionally, CO₂ emission rates were measured to estimate soil respiration activity. Net mineralisation and net nitrification rates were determined by the buried bag method every month. Ion exchange resin bags were used to determine the N availability in the root zone.

Mean N₂O emission rate was 0.9 kg N ha⁻a⁻, which corresponds to 5 % of the N deposited in the forest ecosystem. The main influencing factors were air and soil temperature and NO ⁻₃ accumulated on the ion exchange resin bags. In the course of net ammonification, 14 kg NH ⁺₄ −N ha⁻ were produced per year. About the same amount of NO ⁻₃ −N was formed during nitrification, indicating a rather complete nitrification going on at the site. NO ^t-₃ concentrations found on the ion exchange resin bags were about 3 times as high as NO ^t-₃ produced during net nitrification, indicating substantial NO ^t-₃ immobilisation. The results of this study indicate significant nitrification activities taking place at the Mühleggerköpfl.

     

Variation of stabilised,microbial and biologically active carbon and nitrogen in soil under contrasting land use and agricultural management practices

Land use and agricultural practices modify both the amounts and properties of C and N in soil organic matter. In order to evaluate land use and management-dependent modifications of stable and labile C and N soil pools, (i). organic C and total N content, (ii). microbial (C(mic)) and N (N(mic)) content and (iii). C and N mineralisation rates, termed biologically active C and N, were estimated in arable, grassland and forest soils from northern and southern Germany. The C/N-ratios were calculated for the three levels (i)-(iii) and linked to the eco-physiological quotients of biotic-fixed C and N (C(mic)/C(org), N(mic)/N(t)) and biomass-specific C and N mineralisation rate (qCO(2), qN(min)). Correlations could mainly be determined between organic C, total N, C(mic), N(mic) and C mineralisation for the broader data set of the land use systems. Generally, the mineralisation activity rate at 22 degrees C was highly variable and ranged between 0.11 and 17.67 microg CO(2)-C g(-1) soil h(-1) and -0.12 and 3.81 microg (deltaNH(4)(+)+deltaNO(3)(-))-N g(-1) soil h(-1). Negative N data may be derived from both N immobilisation and N volatilisation during the experiments. The ratio between C and N mineralisation rate differed significantly between the soils ranging from 5 to 37, and was not correlated to the soil C/N ratio and C(mic)/N(mic) ratio. The C/N ratio in the 'biologically active' pool was significantly smaller in soils under conventional farming than those under organic farming systems. In a beech forest, it increased from the L, Of to the Ah horizon. The biologically active C and N pools refer to the current microbial eco-physiology and are related to the need for being C and N use efficient as indicated by metabolic qCO(2) and qN(min) quotients.     

Effects of experimental nitrogen additions on plant diversity in tropical forests of contrasting disturbance regimes in southern China

Responses of understory plant diversity to nitrogen (N) additions were investigated in reforested forests of contrasting disturbance regimes in southern China from 2003 to 2008: disturbed forest (with harvesting of understory vegetation and litter) and rehabilitated forest (without harvesting). Experimental additions of N were administered as the following treatments: Control, 50 kg N ha⁻¹ yr⁻¹, and 100 kg N ha⁻¹ yr⁻¹. Nitrogen additions did not significantly affect understory plant richness, density, and cover in the disturbed forest. Similarly, no significant response was found for canopy closure in this forest. In the rehabilitated forest, species richness and density showed no significant response to N additions; however, understory cover decreased significantly in the N-treated plots, largely a function of a significant increase in canopy closure. Our results suggest that responses of plant diversity to N deposition may vary with different land-use history, and rehabilitated forests may be more sensitive to N deposition.     

Changes in conifer and deciduous forest foliar and forest floor chemistry and basal area tree growth across a nitrogen (N) deposition gradient in the northeastern US

We evaluated foliar and forest floor chemistry across a gradient of N deposition in the Northeast at 11 red spruce (Picea rubens Sarg.) sites in 1987/1988 and foliar and forest floor chemistry and basal area growth at six paired spruce and deciduous sites in 1999. The six red spruce plots were a subset of the original 1987/1988 spruce sites. In 1999, we observed a significant correlation between mean growing season temperature and red spruce basal area growth. Red spruce and deciduous foliar %N correlated significantly with N deposition. Although N deposition has not changed significantly from 1987/1988 to 1999, net nitrification potential decreased significantly at Whiteface. This decrease in net potential nitrification is not consistent with the N saturation hypothesis and suggests that non-N deposition controls, such as climatic factors and immobilization of down dead wood, might have limited N cycling.     

The East African decapod crab Neosarmatium meinerti (de Man) sweeps mangrove floors clean of leaf litter

Leaf litter removal by the abundant mangrove decapod crab Neosarmatium meinerti was studied in series of field and laboratory experiments in East Africa. In the high intertidal Avicennia marina zone crabs buried all leaves placed on the forest floor and consumed on average 67% of them within 2 hrs. High shore crabs in Kenya buried 4 g m(-2) leaf-litter in 1 hr, i.e. approx. twice the daily litter fall. In contrast, in the low shore Sonneratia alba zone, where typical leaf-eating crabs were absent, none of the offered leaves showed signs of herbivory. Leaf choice experiments in the laboratory showed that N. meinerti preferred some species to others. Leaf consumption per gram crab was higher in females than males. The laboratory studies also indicated that crabs could consume substantially more than the average daily litter fall. Video recordings documented frequent fights to gain or retain fallen leaves, suggesting strong competition for leaf litter. Earlier studies indicating that N. meinerti may sweep mangrove forest floors clean of leaf litter are confirmed. In high shore mangroves of East and South Africa where N. meinerti is common, energy flow appears unique: virtually all litter production is retained.     

Interactive effects of CO2 and O3 on a ponderosa pine plant/litter/soil mesocosm.

To study individual and combined impacts of two important atmospheric trace gases, CO2 and O3, on C and N cycling in forest ecosystems; a multi-year experiment using a small-scale ponderosa pine (Pinus ponderosa Laws.) seedling/soil/litter system was initiated in April 1998. The experiment was conducted in outdoor, sun-lit chambers where aboveground and belowground ecological processes could be studied in detail. This paper describes the approach and methodology used, and presents preliminary data for the first two growing seasons. CO2 treatments were ambient and elevated (ambient + 280 ppm). O3 treatments were elevated (hourly averages to 159 ppb, cumulative exposure > 60 ppb O3, SUM 06 approximately 10.37 ppm h), and a low control level (nearly all hourly averages <40 ppb. SUM 06 approximately 0.07 ppm h). Significant (P < 0.05) individual and interactive effects occurred with elevated CO2 and elevated O3. Elevated CO2 increased needle-level net photosynthetic rates over both seasons. Following the first season, the highest photosynthetic rates were for trees which had previously received elevated O3 in addition to elevated CO2. Elevated CO2 increased seedling stem diameters, with the greatest increase at low O3. Elevated CO2 decreased current year needle % N in the summer. For 1-year-old needles measured in the fall there was a decrease in % N with elevated CO2 at low O3, but an increase in % N with elevated CO2 at elevated O3. Nitrogen fixation (measured by acetylene reduction) was low in ponderosa pine litter and there were no significant CO2 or O3 effects. Neither elevated CO2 nor elevated O3 affected standing root biomass or root length density. Elevated O3 decreased the % N in coarse-fine (1-2 mm diameter) but not in fine (< 1 mm diameter) roots. Both elevated CO2 and elevated O3 tended to increase the number of fungal colony forming units (CFUs) in the AC soil horizon, and elevated O3 tended to decrease bacterial CFUs in the C soil horizon. Thus, after two growing seasons we showed interactive effects of O3 and CO2 in combination, in addition to responses to CO2 or O3 alone for a ponderosa pine plant/litter/soil system.     

Detrital control on the release of dissolved organic nitrogen (DON) and dissolved inorganic nitrogen (DIN) from the forest floor under chronic N deposition

The role of detrital quantity and quality in forest floor N leaching was investigated in a litter manipulation experiment at a deciduous forest under chronic N deposition. Dissolved inorganic nitrogen (DIN) comprised the bulk of nitrogen leaching from the control except a short period following autumn litterfall. The dominance of DIN was strengthened by litter exclusion, whereas the addition of glucose or fresh litter led to a small increase in dissolved organic nitrogen (DON) and either a temporary or gradual reduction in NO(3)(-) release, respectively. Changes in soluble organic C and microbial C in the forest floor implied that increased availability of C sources might have enhanced microbial immobilization of DIN, either temporarily following glucose application or over the longer term following litter addition. The results suggest that detrital quantity and quality can play a crucial role in determining the balance between DIN and DON in N-enriched forest soils.     

Regional patterns in foliar (15)N across a gradient of nitrogen deposition in the northeastern US

Recent studies have demonstrated that natural abundance (15)N can be a useful tool for assessing nitrogen saturation, because as nitrification and nitrate loss increase, delta(15)N of foliage and soil also increases. We measured foliar delta(15)N at 11 high-elevation spruce-fir stands along an N deposition gradient in 1987-1988 and at seven paired northern hardwood and spruce-fir stands in 1999. In 1999, foliar delta(15)N increased from -5.2 to -0.7 per thousand with increasing N deposition from Maine to NY. Foliar delta(15)N decreased between 1987-1988 and 1999, while foliar %N increased and foliar C:N decreased at most sites. Foliar delta(15)N was strongly correlated with N deposition, and was also positively correlated with net nitrification potential and negatively correlated with soil C:N ratio. Although the increase in foliar %N is consistent with a progression towards N saturation, other results of this study suggest that, in 1999, these stands were further from N saturation than in 1987-1988.     

Diversity of shrub tree layer, leaf litter decomposition and N release in a Brazilian Cerrado under N, P and N plus P additions

This study investigated changes in diversity of shrub-tree layer, leaf decomposition rates, nutrient release and soil NO fluxes of a Brazilian savanna (cerrado sensu stricto) under N, P and N plus P additions. Simultaneous addition of N and P affected density, dominance, richness and diversity patterns more significantly than addition of N or P separately. Leaf litter decomposition rates increased in P and NP plots but did not differ in N plots in comparison to control plots. N addition increased N mass loss, while the combined addition of N and P resulted in an immobilization of N in leaf litter. Soil NO emissions were also higher when N was applied without P. The results indicate that if the availability of P is not increased proportionally to the availability of N, the losses of N are intensified.     

Soil functional responses to excess nitrogen inputs at global scale

There is little evidence that nitrogen (N) cycling in the highly weathered, low-phosphorus (P), acidic soils found in Southern Hemisphere continents will differ greatly from that in North America and Europe. Evidence from the 'south' shows: the similarity in forms and temporal patterns in losses of N from different land uses; that the C:N ratios of the forest floor/litter layer from different continents are strongly predictive of a range of processes on a global scale; that generalizations based on Northern Hemisphere experience of the impact of N additions to 'P-limited' ecosystems are likely to fail for southern ecosystems where anatomical and physiological adaptation of native plants to low-P soils makes questionable the concept of 'P-limitation'; that the greatest threats in the 'south' are probably changes in land use that may greatly increase N inputs and turnover; that localized increases in N inputs produce similar effects to those seen in the 'north'.     

Combined effects of nitrogen addition and organic matter manipulation on soil respiration in a Chinese pine forest

The response of soil respiration (Rs) to nitrogen (N) addition is one of the uncertainties in modelling ecosystem carbon (C). We reported on a long-term nitrogen (N) addition experiment using urea (CO(NH₂)₂) fertilizer in which Rs was continuously measured after N addition during the growing season in a Chinese pine forest. Four levels of N addition, i.e. no added N (N0: 0 g N m⁻² year⁻¹), low-N (N1: 5 g N m⁻² year⁻¹), medium-N (N2: 10 g N m⁻² year⁻¹), and high-N (N3: 15 g N m⁻² year⁻¹), and three organic matter treatments, i.e. both aboveground litter and belowground root removal (LRE), only aboveground litter removal (LE), and intact soil (CK), were examined. The Rs was measured continuously for 3 days following each N addition application and was measured approximately 3–5 times during the rest of each month from July to October 2012. N addition inhibited microbial heterotrophic respiration by suppressing soil microbial biomass, but stimulated root respiration and CO₂ release from litter decomposition by increasing either root biomass or microbial biomass. When litter and/or root were removed, the “priming” effect of N addition on the Rs disappeared more quickly than intact soil. This is likely to provide a point of view for why Rs varies so much in response to exogenous N and also has implications for future determination of sampling interval of Rs measurement.

     

Spatial and temporal small-scale variability of nitrogen mobilization in a forest ecosystem with high N deposition in NW-Germany

For conifer stands in NW-Germany with high DIN load (23-35 kg N ha⁻¹ a⁻¹) and a long history of nitrogen export the risk of N mobilization were investigated. Ammonium is the most mobilized N species, pointing towards either conditions not favoring nitrification or, more likely - under the dominant aerobic conditions - a very high amount of ammonium in the forest floor. Independence of net nitrification and net ammonification from each other indicates the existence of two separate systems. The nitrifying system depends very much on biotic conditions - as a function of energy and moisture - and seems not to be directly related to N deposition. In contrast, for the ammonification system (Oe horizon) a correlation with the sum of ammonium deposition three months prior to sampling was found. However, the role of disturbance, i.e. nitrogen export, during the last centuries and the role of recovery of the N balance during the last 150 years is still not clear.     

Biomineralisation of 1,2,4-trichlorobenzene in soils by an adapted microbial population

In laboratory experiments the mineralisation of 14C-labelled 1,2,4-trichlorobenzene (1,2,4-TCB) in soils was studied by direct measurement of the evolved 14CO2. The degradation capacity of the indigenous microbial population was investigated in an agricultural soil and in a soil from a contaminated site. Very low mineralisation of 1% within 23 days was measured in the agricultural soil. Whereas in the soil from the contaminated site the mineralisation occurred very fast and in high rates; up to 62% of the initially applied amount of 1,2,4-TCB were mineralised within 23 days. The transfer of the adapted microbial population into the agricultural soil significantly enhanced the mineralisation of 1,2,4-TCB in this soil, reflecting, that the transferred microbial population survived and maintained its degradation ability in the new microbial ecosystem. Additional nutrition sources ((NH4)2HPO4) increased the mineralisation rates in the first days significantly in the contaminated soil. In the soil from the contaminated site high amounts of non extractable 14C-residues were formed.     

Effects of increased deposition of atmospheric nitrogen on an upland moor: nitrogen budgets and nutrient accumulation

This study was designed to investigate the effect of long-term (11 years) ammonium nitrate additions on standing mass, nutrient content (% and kg ha(-1)), and the proportion of the added N retained within the different compartments of the system. The results showed that more than 90% of all N in the system was found in the soil, particularly in the organic (Oh) horizon. Added N increased the standing mass of vegetation and litter and the N content (kg N ha(-1)) of almost all measured plant, litter and soil compartments. Green tissue P and K content (kg ha(-1)) were increased, and N:P ratios were increased to levels indicative of P limitation. At the lowest treatment, most of the additional N was found in plant/litter compartments, but at higher treatments, there were steep increases in the amount of additional N in the underlying organic and mineral (Eag) horizons. The budget revealed that the proportion of added N found in the system as a whole increased from 60%, 80% and up to 90% in response to the 40, 80 and 120 kg N ha(-1) year(-1) treatments, respectively.     

Nitrogen biogeochemistry in the Adirondack Mountains of New York: hardwood ecosystems and associated surface waters

Studies on the nitrogen (N) biogeochemistry in Adirondack northern hardwood ecosystems were summarized. Specific focus was placed on results at the Huntington Forest (HFS), Pancake-Hall Creek (PHC), Woods Lake (WL), Ampersand (AMO), Catlin Lake (CLO) and Hennessy Mountain (HM). Nitrogen deposition generally decreased from west to east in the Adirondacks, and there have been no marked temporal changes in N deposition from 1978 through 1998. Second-growth western sites (WL, PHC) had higher soil solution NO(3-) concentrations and fluxes than the HFS site in the central Adirondacks. Of the two old-growth sites (AMO and CLO), AMO had substantially higher NO(3-) concentrations due to the relative dominance of sugar maple that produced litter with high N mineralization and nitrification rates. The importance of vegetation in affecting N losses was also shown for N-fixing alders in wetlands. The Adirondack Manipulation and Modeling Project (AMMP) included separate experimental N additions of (NH4)2SO4 at WL, PHC and HFS and HNO3 at WL and HFS. Patterns of N loss varied with site and form of N addition and most of the N input was retained. For 16 lake/watersheds no consistent changes in NO(3-) concentrations were found from 1982 to 1997. Simulations suggested that marked NO(3-) loss will only be manifested over extended periods. Studies at the Arbutus Watershed provided information on the role of biogeochemical and hydrological factors in affecting the spatial and temporal patterns of NO(3-) concentrations. The heterogeneous topography in the Adirondacks has generated diverse landscape features and patterns of connectivity that are especially important in regulating the temporal and spatial patterns of NO(3-) concentrations in surface waters.     

Distribution and movement of nutrients and metals in a Pinus radiata forest soil following applications of biosolids

Samples of biosolids, spiked with increasing amounts of Cu, Ni or Zn were applied to field plots in a Pinus radiata forest, and the nutrient and metal status of the forest litter and underlying mineral soil was monitored over a period of six years following application. The macronutrient status of the forest litter was changed markedly by the biosolids application, with substantial increases in N, P and Ca concentrations, and decreases in Mg and K. The C/N ratio of the litter was also decreased and pH was increased by the biosolids application. The metals applied with the biosolids were retained predominantly in the litter layer, and even with non-metal-spiked biosolids there were substantial increases in litter metal concentrations. There was also firm evidence of some movement of Cu, Ni and Zn into the underlying mineral soil. The potential environmental issues resulting from these changes in nutrient and metal status are discussed.     

Tracking nitrogen losses in a greenhouse crop rotation experiment in North China using the EU-Rotate_N simulation model

Vegetable production in China is associated with high inputs of nitrogen, posing a risk of losses to the environment. Organic matter mineralisation is a considerable source of nitrogen (N) which is hard to quantify. In a two-year greenhouse cucumber experiment with different N treatments in North China, non-observed pathways of the N cycle were estimated using the EU-Rotate_N simulation model. EU-Rotate_N was calibrated against crop dry matter and soil moisture data to predict crop N uptake, soil mineral N contents, N mineralisation and N loss. Crop N uptake (Modelling Efficiencies (ME) between 0.80 and 0.92) and soil mineral N contents in different soil layers (ME between 0.24 and 0.74) were satisfactorily simulated by the model for all N treatments except for the traditional N management. The model predicted high N mineralisation rates and N leaching losses, suggesting that previously published estimates of N leaching for these production systems strongly underestimated the mineralisation of N from organic matter.     

Impact of crude oil exposure on nitrogen cycling in a previously impacted Juncus roemerianus salt marsh in the northern Gulf of Mexico

This study investigated potential nitrogen fixation, net nitrification, and denitrification responses to short-term crude oil exposure that simulated oil exposure in Juncus roemerianus salt marsh sediments previously impacted following the Deepwater Horizon accident. Temperature as well as crude oil amount and type affected the nitrogen cycling rates. Total nitrogen fixation rates increased 44 and 194 % at 30 °C in 4,000 mg kg^?1 tar ball and 10,000 mg kg^?1 moderately weathered crude oil treatments, respectively; however, there was no difference from the controls at 10 and 20 °C. Net nitrification rates showed production at 20 °C and consumption at 10 and 30 °C in all oil treatments and controls. Potential denitrification rates were higher than controls in the 10 and 30?ºC treatments but responded differently to the oil type and amount. The highest rates of potential denitrification (12.7?±?1.0 nmol N g^?1 wet h^?1) were observed in the highly weathered 4,000 mg kg^?1 oil treatment at 30 °C, suggesting increased rates of denitrification during the warmer summer months. These results indicate that the impacts on nitrogen cycling from a recurring oil spill could depend on the time of the year as well as the amount and type of oil contaminating the marsh. The study provides evidence for impact on nitrogen cycling in coastal marshes that are vulnerable to repeated hydrocarbon exposure.     

Urbanization-related changes in European aspen (Populus tremula L.): Leaf traits and litter decomposition

We investigated foliar and litter responses of European aspen (Populus tremula L.) to urbanization, including factors such as increased temperature, moisture stress and nitrogen (N) deposition. Leaf samples were collected in 2006-2008 from three urban and three rural forest stands in the Helsinki Metropolitan Area, southern Finland, and reciprocal litter transplantations were established between urban and rural sites. Urban leaves exhibited a higher amount of epicuticular waxes and N concentration, and a lower C:N ratio than rural ones, but there was no difference in specific leaf area. Urban litter had a slightly higher N concentration, lower concentrations of lignin and total phenolics, and was more palatable to a macrofaunal decomposer. Moreover, litter decay was faster at the urban site and for urban litter. Urbanization thus resulted in foliar acclimatization in terms of increased amount of epicuticular waxes, as well as in accelerated decomposition of the N-richer leaf litter.