The role of monoterpenes in soil nitrogen cycling processes in ponderosa pine

The effects of select monoterpenes on nitrogen (N) mineralization and nitrification potentials were determined in four separate laboratory bioassays. The effect of increasing monoterpene addition was an initial reduction in NO₃^–-N production (nitrification inhibition), followed by a reduction in the sum of NH₄⁺-N and NO₃^–-N (inhibition of net N mineralization and net immobilization at high monoterpene additions. Monoterpenes could produce this pattern by inhibiting nitrification, reducing net N mineralization, enhancing immobilization of NO₃^–-N relative to NH₄⁺-N, and/or stimulating overall net immobilization of N by carbon-rich material.Initial monoterpene concentrations in the assay soils were about 5% of the added amount and were below detection after incubation in most samples.Potential N mineralization-immobilization, nitrification, and soil monoterpene concentrations were determined by soil horizon for four collections from a ponderosa pine (Pinus ponderosa) stand in New Mexico. Concentrations of monoterpenes declined exponentially with soil depth and varied greatly within a horizon. Monoterpene content of the forest floor was not correlated with forest floor biomass. Net N mineralization was inversely correlated with total monoterpene content of all sampled horizons. Nitrification was greatest in the mineral soil, intermediate in the F-H horizon, and never occurred in the L horizon. Nitrification in the mineral soil was inversely correlated with the amount of monoterpenes in the L horizon that contain terminal unsaturated carbon-carbon bonds (r² = 0.37, P 0.01). This pattern in the field corresponded to the pattern shown in the laboratory assays with increasing monoterpene additions.     

Investigation of nitrification in forest soils with soil percolation columns

Soil percolation columns in which a pF of 2 could be maintained were developed to study nitrification in soils and litter of an acid and a calcareous forest soil location. High nitrification rates were observed in the calcareous soil. In the acid soil nitrification was much slower. A column filled with leaf litter gave a low nitrification rate at the start of the experiment, but a high rate was found after 60 days of percolation with an ammonium-containing medium of pH 4. In this leaf litter high numbers of autotrophic bacteria were just present at the beginning of the experiment, whereas at the end only low numbers were detected. Results indicate that autotrophic bacteria from acid soils are sensitive to a pH increase.     

Loss of nitrogen in compacted grassland soil by simultaneous nitrification and denitrification

The soils of mid-Wales in grazed permanent pasture usually exhibit stagnogley features in the top 4–10 cm even though on sloping sites, they are freely drained. Nitrogen is often poorly recovered under these conditions. Our previous studies suggest that continuing loss of available N through concurrent nitrification and denitrification might provide an explanation for poor response to fertilizer N. The work described was designated to further test this proposition. When NH ₄ ⁺ –N was applied to the surface of intact cores, equilibrated at –5kPa matric potential, about 70% of NH ₄ ⁺ –N initially present was lost within 56 days of incubation. Study of different sections of the cores showed a rise in NO ₃ ^- level in the surface 0–2.5 cm soil layer but no significant changes below this depth. The imbalance between NO ₃ ^- accumulation and NH ₄ ⁺ disappearance during the study indicated a simultaneous nitrification and denitrification in the system. Furthermore, the denitrification potential of the soil was 3–4 times greater than nitrification potential so no major build-up of NO ₃ ^- would be expected when two processes occur simultaneously in micro-scale. When nitrification was inhibited by nitrapyrin, a substantial amount of NH ₄ ⁺ –N remained in the soil and persisted till the end of the incubation. The apparent recovery of applied N increased and of the total amount of N applied, 50% more was recovered relative to without nitrapyrin. It appears that addition of nitrapyrin inhibited nitrification, and consequently denitrification, by limiting the supply of NO ₃ ^- for denitrifying organisms. Emission of N₂O from the NH ₄ ⁺ amended soil cores further confirmed that loss of applied N was the result of both nitrification and denitrification, which occurred simultaneously in adjacent sites at shallow depths. This N loss could account for the poor response to fertilizer N often observed in pastoral agriculture in western areas of the UK.     

Decline of soil nitrogen mineralization and nitrification during forest conversion of evergreen broad-leaved forest to plantations in the subtropical area of Eastern China

We examined soil nitrogen (N) mineralization and nitrification rates, and soil and forest floor properties in one native forest: evergreen broad-leaved forest (EBLF), one secondary shrubs (SS), and three adjacent plantation forests: Chinese fir plantation (CFP), bamboo plantation (BP) and waxberry groves (WG) in Tiantong National Forest Park, Eastern China. All forests showed seasonal dynamics of N mineralization and nitrification rates. Soil N mineralization rate was highest in EBLF (1.6 ± 0.3 mg-N kg⁻¹ yr⁻¹) and lowest in CFP (0.4 ± 0.1 mg-N kg⁻¹ yr⁻¹). Soil nitrification rate was also highest in EBLF (0.6 ± 0.1 mg-N kg⁻¹ yr⁻¹), but lowest in SS (0.02 ± 0.01 mg-N kg⁻¹ yr⁻¹). During forest conversion of EBLF to SS, CFP, BP and WG, soil N mineralization rate (10.7%, 73%, 40.3% and 69.8%, respectively), soil nitrification rate (94.9%, 32.2%, 33.9% and 39%, respectively), and soil N concentration (50%, 65.4%, 78.9% and 51.9%, respectively) declined significantly. Annual soil N mineralization was positively correlated with total C and N concentrations of surface soil and total N concentration of forest floor, and negatively correlated with soil bulk density, soil pH and C:N ratio of forest floor across the five forests. Annual soil nitrification was positively correlated with total C concentration of surface soil and N concentration of forest floor, and negatively correlated with soil bulk density and forest floor mass. In contrast, annual soil nitrification was not correlated to pH value, total N concentration, C:N ratio of surface soil and total C concentration and C:N ratio of forest floor.     

Seasonal and annual variation in nitrogen mineralization and nitrification along an elevational gradient in New Mexico

Patterns and amounts of nitrogen loss from disturbed ecosystems vary widely. The mineralization of organic nitrogen to ammonium and then nitrification to nitrate are important processes regulating nitrogen cycling rates and nitrogen losses. Nitrification is a significant process because of the production of the nitrate anion which is easily leached or denitrified. Most studies of these processes do not evaluate their seasonal and yearly variations. This study demonstrates that marked seasonal and yearly variations can occur in these processes in different ecosystems and suggests that nitrogen loss or other system properties correlated with one arbitrarily selected collection can be misleading. Spruce-fir and ponderosa pine ecosystems demonstrated little actual orpotential nitrification. Aspen and mixed conifer ecosystems demonstrated distinct seasonal patterns with increased rates of mineralization and nitrification during spring and summer months and a precipitous decline in both rates coincident with autumn foliage litterfall.The relative availability of soil nitrogen along with the amount of nitrogen circulating annually in litterfall prior to disturbance are useful predictors of the potential for nitrate production and loss following disturbance. However, other controls, including regulation by organic compounds, appear important in determining seasonal and annual variation in actual nitrification rates.     

植物挥发性单萜经土壤载体的化感作用——以三裂叶豚草(Ambrosia trifida L.)为例

植物释放的挥发性单萜在生态系统中起着重要的作用,这些单萜不仅能以空气为载体对其他植物显示直接的化感作用,而且也能以土壤为载体进行间接的化感作用.通过对三裂叶豚草挥发物对植物种子萌发和土壤微生物种群影响及其化学成分的鉴定研究,验证了以单萜类物质为主的三裂叶豚草挥发物可以经土壤载体对其他植物及土壤微生物显示化感效应.还对植物挥发物化学成分鉴定的GC(气相色谱)和GC-MS(气相色谱-质谱联用)方法存在的一些误区进行了澄清,这将有助于对生态系统中植物挥发物化感作用的研究和认识.     

Influence of tree species and ground vegetation on nitrification in an acid forest soil

Soil N transformations were studied at Ironhill, near Liphook, UK as part of a forest fumigation experiment. Nitrification potential was measured in a humoferric podzol soil, of pH 3 (in 0.01 M CaCl₂). An initial experiment into nitrogen mineralisation potential indicated that nitrification was linked strongly to the species of coniferous tree growing in the soil. Transfer of soil solution between soils had no influence on mineralisation potential and allelopathic effects of the trees were not demonstrated. The initial finding was attributed subsequently to the type of ground vegetation and its management. Attempts to reproduce soil conditions, which promoted nitrification, were partially successful.Soil, from the Ironhill site, was incubated with various nitrogenous substrates and other nutrients and sources of carbon to test whether heterotrophs were responsible for nitrification. Organic N (which was ammonified) promoted nitrification, but the addition of ammonium was inhibitory unless supplied with a readily available carbon source such as acetate. Nitrification potential was unaffected when soils were incubated with an inhibitor of autotrophic nitrification. The results of these experiments supported strongly the hypothesis that heterotrophic organisms were responsible for nitrification in this soil.     

Temperature-dependent nitrogen transformations in acid oak-beech forest litter in the Netherlands

Laboratory incubation experiments have been carried out to quantify net nitrogen mineralization and nitrification in oak-beech litter at temperatures ranging from 0 to 30°C. Net mineralization was linearly proportional to temperature. Nitrification was inhibited at 0,5 and 30°C. As compared with soils under cultivation, there is only restricted knowledge of nitrification kinetics in acid forest litters, especially when temperature is considered. With these litter types, one should be cautious applying high incubation temperatures, which seldomly occur under field conditions.     

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO₂. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage. The major contributors to soil‐surface CO₂ effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large‐scale girdling experiment was performed in a long‐term nutrient optimization experiment in a 40‐year‐old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil‐surface CO₂ fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots. In late July, the time of the seasonal maximum in soil‐surface CO₂ efflux, the total soil‐CO₂ efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil‐surface CO₂ efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated. Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.     

The relative importance of autotrophic and heterotrophic nitrification in a conifer forest soil as measured by 15N tracer and pool dilution techniques

The importance of heterotrophic nitrification was studied in soil from a mixed-conifer forest. Three sites in the forest were sampled: a clear cut area, a young stand and a mature stand. In the mature stand, the mineral soil (0–10 cm) and the organic layer were sampled separately. Gross rates of N mineralization and nitrification were measured by¹⁵NH ₄ ⁺ and¹⁵NO ₃ ^– isotopic pool dilution, respectively. The rates of autotrophic and heterotrophic nitrification were distinguished by use of acetylene as a specific inhibitor of autotrophic nitrification. In samples supplemented with¹⁵NH ₄ ⁺ and treated with acetylene, no¹⁵NO ₃ ^– was detectable showing that the acetylene treatment effectively blocked the autotrophic nitrification, and that NH ₄ ⁺ was not a substrate for heterotrophic nitrification. In the clear cut area, autotrophic nitrification was the most important NO ₃ ^– generating process with total nitrification (45 ug N kg^–1h^–1) accounting for about one-third of gross N mineralization (140 ug N kg^–1 h^–1). In the young and mature forested sites, gross nitrification rates were largely unaffected by acetylene treatment indicating that heterotrophic nitrification dominated the NO ₃ ^– generating process in these areas. In the mature forest mineral and organic soil, nitrification (heterotrophic) was equal to only about 5% of gross mineralization (gross mineralization rates of 90 ug N kg^–1 h^–1 mineral; 550 ug N kg^–1 h^–1 organic). The gross nitrification rate decreased from the clear cut area to the young forest area to the mineral soil of the mature forest (45; 17; 4.5 ug kg^–1 h^–1 respectively). The¹⁵N isotope pool dilution method, combined with acetylene as an inhibitor of autotrophic nitrification provided an effective technique for assessing the importance of heterotrophic nitrification in the N-cycle of this mixed-conifer ecosystem.     

Nitrogen cycling and proton fluxes in an acid forest soil

Ironhill, near Liphook, UK, was the site of a forest fumigation experiment. Nitrogen cycling within the humoferric podzol soil was a component of the study into the impacts of sulphur dioxide and ozone on coniferous trees. Variation in total soil N and N mineralisation was too great to determine impacts from the fumigant gases. Differences in the nitrogen mineralisation potential of the soils were unrelated to the initial levels of mineral or total N, or to pH. Mineralisation potential was affected by temperature and a Q₁₀ of approximately 3 was demonstrated. Mineralisation potential was reduced in very dry soils, but the wetting of these dry soils did not result in enhanced mineralisation, relative to fresh samples of equivalent moisture content. Nitrification potential was detected in this forest soil of pH 3 (in 0.01 m CaCl₂).The soil N data and those from the analysis of N within vegetation were used to prepare N budgets for the second and third seasons' growth of a mixed conifer forest; by the third year, N appeared to limit tree growth.The relative magnitude of proton fluxes from plant growth, nitrification and atmospheric inputs was estimated. Acidity generated from the balance of cations and anions in plant uptake, and soil N transformations was estimated to be comparable to that from `acid rain'. This comparison was based on only parts of the N cycle because they may occur remotely, in time or space, from other transformations of N. The comparison is valid, therefore, at the scale of individual trees or small-scale experimental plots, but at forest scale, wet and dry deposition were predicted to be the more significant for ecosystem acidification.     

Soil N mineralization and nitrification in relation to nitrogen solution chemistry in a small forested watershed

Spatial variations in soil processes regulating mineral N losses to streams were studied in a small watershed near Toronto, Ontario. Annual net N mineralization in the 0–8 cm soil was measured in adjacent upland and riparian forest stands using in situ soil incubations from April 1985 to 1987. Mean annual rates of soil N mineralization and nitrification were higher in a maple soil (93.8 and 87.0 kg.ha^–1) than in a pine soil (23.3 and 8.2 kg.ha^–1 ). Very low mean rates of mineralization (3.3 kg.ha^–1) and nitrification (3.4 kg.ha^–1) were found in a riparian hemlock stand. Average NO₃-N concentrations in soil solutions were 0.3–1.0 mg.L^–1 in the maple stand and >0.06mg.L^–1 in the pine stand. Concentrations of NO₃–N in shallow ground water and stream water were 3–4× greater in a maple subwatershed than in a pine subwatershed. Rapid N uptake by vegetation was an important mechanism reducing solution losses of NO₃–N in the maple stand. Low rates of nitrification were mainly responsible for negligible NO₃–N solution losses in the pine stand.     

The use of different soil nitrogen sources by young Norway spruce plants

 Three-year-old Norway spruce trees were planted into a low-nitrogen mineral forest soil and supplied either with two different levels of mineral nitrogen (NH₄NO₃) or with a slow-release form of organic nitrogen (keratin). Supply of mineral nitrogen increased the concentrations of ammonium and nitrate in the soil solution and in CaCl₂-extracts of the rhizosphere and bulk soil. In the soil solution, in all treatments nitrate concentrations were higher than ammonium concentrations, while in the soil extracts ammonium concentrations were often higher than nitrate concentrations. After 7 months of growth, ¹⁵N labelled ammonium or nitrate was added to the soil. Plants were harvested 2 weeks later. Keratin supply to the soil did not affect growth and nitrogen accumulation of the trees. In contrast, supply of mineral nitrogen increased shoot growth and increased the ratio of above-ground to below-ground growth. The proportion of needle biomass to total above-ground biomass was not increased by mineral N supply. The atom-% ¹⁵N was higher in younger needles than in older needles, and in younger needles higher in plants supplied with ¹⁵N-nitrate than in plants supplied with ¹⁵N-ammonium. The present data show that young Norway spruce plants take up nitrate even under conditions of high plant internal N levels. Received: 1 April 1998 / Accepted: 9 October 1998     

Net nitrogen mineralization,nitrification and CO2 production in alternating moisture conditions in an unfertilized low-humus sandy soil from the Sahel

In this study the rates of net mineralization, net immobilization and net nitrification have been quantified under laboratory conditions in a sandy low-humus soil from a semi-arid region, in absence of plant growth. Incubation experiments were carried out under constant humidity and under alternating wet and dry conditions to simulate field conditions during the rainy season. The ammonium and nitrate content of the incubates were determined and their CO₂ production measured.The rate of net mineralization at field capacity was 0.6 kg N ha^–1d^–1 during the first 40 days and decreased to 0.06 kg N ha^–1d^–1 after 400 days. This rate was twice as high on wet days under alternating wet and dry conditions. The rate of net nitrification during alternating wet and dry conditions was also higher (1.9 kg N ha^–1d^–1) than at constant field capacity (1.3 kg N ha^–1d^–1) until the ammonium was almost completely depleted. These rates of net mineralization and net nitrification are in agreement with field observations.Net immobilization did not occur in the experiments, unless glucose was added to the soil.The data on CO₂ production and net mineralization showed that the C/N ratio of the degraded material was around 9 or below. It is much lower than the ratio of total carbon over total nitrogen in the soil. This indicates that microorganisms and compounds high in nitrogen were mineralized. Certainly after about 30 days the only growth taking place is based on turnover of material of the microbial biomass itself.A decrease in the amount of inorganic nitrogen was observed upon drying of the soil, while it returned to the original content after rewetting. It is postulated that this might be due to temporary uptake of nitrogen in an inorganic form in microorganisms as part of their osmoregulation.     

Grass species and soil type effects on microbial biomass and activity

We evaluated plant versus soil type controls on microbial biomass and activity by comparing microbial biomass C, soil respiration, denitrification potential, potential net N mineralization and nitrification in different soils supporting four grass species, and by growing a group of 10 different grass species on the same soil, in two experiments respectively. In the first experiment, none of the microbial variables showed significant variation with grass species while all variables showed significant variation with soil type, likely due to variation in soil texture. In the second experiment, there were few significant differences in microbial biomass C among the 10 grasses but there were significant relationships between variation in microbial biomass C and potential net N mineralization (negative), soil respiration (positive) and denitrification (positive). There was no relationship between microbial biomass C and either plant yield or plant N concentration. The results suggest that 1) soil type is a more important controller of microbial biomass and activity than grass species, 2) that different grass species can create significant, but small and infrequent, differences in microbial biomass and activity in soil, and 3) that plant-induced variation in microbial biomass and activity is caused by variation in labile C input to soil.     

Short-term increases in relative nitrification rates due to trenching in forest floor and mineral soil horizons of different forest types

The representation of NO₃ ^– dynamics within forest growth simulation models could improve forest management. An extensive literature review revealed an 88% probability of measuring a higher relative nitrification index (i.e. RNI = [NO₃ ^–] ÷ [NO₃ ^– + NH₄ ⁺]) in mineral soil horizons than in forest floors, across a wide range of conifer and hardwood ecosystems. We then hypothesised that humus form and fine root density could be used as two crude variables to predict changes in in situ, potential and relative nitrification rates. Twenty-seven trench plots were established in 1999, across nine contrasting hardwood and coniferous stands in the Eastern Townships of Québec. Forest floor and mineral soil samples were collected from each plot, and from a 1 m radius surrounding each plot, on three dates during summer 2000. In situRNI values increased significantly in trench plots as the season progressed. Potential nitrification rates (i.e. NO₃ ^– concentrations following incubation) were two orders of magnitude higher in forest floor than in mineral soil samples. RNI was significantly higher in mineral soil than in forest floor samples after incubations, but the relative increase in RNI due to trenching was higher in forest floor samples. Indices of available C were significantly higher in forest floor than mineral soil samples, and decreased only in forest floor samples during incubations. Likewise, trenching significantly reduced available C in forest floor samples only. Seasonal changes in soil temperature and fine root growth were the most plausible explanations for seasonal changes in NO₃ ^– dynamics, whereas other factors such soil acidity and moisture appeared less important in determining NO₃ ^– dynamics in this study. We conclude that crude assessments of humus form and fine root density have the potential to be used as calibration parameters for the simulation of NO₃ ^– dynamics in forest growth and yield models.     

Host and non-host odour signals governing host selection by the pine shoot beetle, Tomicus piniperda and the spruce bark beetle, Hylurgops palliatus (Col., Scolytidae)

Abstract:  Tomicus piniperda and Hylurgops palliatus colonize susceptible host trees by responding to host-specific odour signals as well as by avoiding volatiles emanating from non-host conifers. In the field, the pine shoot beetle, T. piniperda , responded in high numbers to natural odour sources provided by their host tree, Pinus sylvestris , while the non-host conifers Larix decidua , Picea abies , or Pseudotsuga menziesii were significantly less attractive. In contrast, the spruce bark beetle, Hylurgops palliatus , preferentially responded to its main host, P. abies . Furthermore, T. piniperda attacks on P. sylvestris bolts decreased in presence of bark and wood particles from the non-host P. abies , whereas particles from P. menziesii appeared not to affect T. piniperda attacks. Apparently, tree-specific volatiles act at close range as specific signals that lead to the successful discrimination and colonization of the respective host tree species.     

Geostatistical analysis of soil properties in a secondary tropical dry forest,St. Lucia,West Indies

Spatial variability of soil properties directly influences forest growth. However, spatial variation in soil properties has not been studied within tropical dry forests. As such, it is unclear whether soil properties, like moisture and N availability, display spatial variation at scales similar to that of other ecosystems. To gain insight into this variation, we established a 56 × 56 m sampling grid in tropical dry forest on the Caribbean island of St. Lucia. Samples collected at 4-m intervals were analyzed for forest floor mass, soil texture, pH, organic C, net N mineralization, net nitrification and available P. Geostatistical procedures were used to determine spatial autocorrelation of the aforementioned properties and processes. Semivariogram parameters were used in a block kriging procedure to produce spatial maps of soil properties. At the scale of our study, most soil properties exhibited spatial autocorrelation at distances of 24 m or less. Varying degrees of similarity were found between patterns of forest floor mass, organic C, net N mineralization, net nitrification and available P. No similarity was found between soil texture or pH and other properties. Fine-scale spatial patterns of net N mineralization and net nitrification are likely driven by overstory litter inputs, rather than variation in soil texture and water availability.