Tree species control rates of free-living nitrogen fixation in a tropical rain forest

Tropical rain forests represent some of the most diverse ecosystems on earth, yet mechanistic links between tree species identity and ecosystem function in these forests remains poorly understood. Here, using free-living nitrogen (N) fixation as a model, we explore the idea that interspecies variation in canopy nutrient concentrations may drive significant local-scale variation in biogeochemical processes. Biological N fixation is the largest "natural" source of newly available N to terrestrial ecosystems, and estimates suggest the highest such inputs occur in tropical ecosystems. While patterns of and controls over N fixation in these systems remain poorly known, the data we do have suggest that chemical differences among tree species canopies could affect free-living N fixation rates. In a diverse lowland rain forest in Costa Rica, we established a series of vertical, canopy-to-soil profiles for six common canopy tree species, and we measured free-living N fixation rates and multiple aspects of chemistry of live canopy leaves, senesced canopy leaves, bulk leaf litter, and soil for eight individuals of each tree species. Free-living N fixation rates varied significantly among tree species for all four components, and independent of species identity, rates of N fixation ranged by orders of magnitude along the vertical profile. Our data suggest that variations in phosphorus (P) concentration drove a significant fraction of the observed species-specific variation in free-living N fixation rates within each layer of the vertical profile. Furthermore, our data suggest significant links between canopy and forest floor nutrient concentrations; canopy P was correlated with bulk leaf litter P below individual tree crowns. Thus, canopy chemistry may affect a suite of ecosystem processes not only within the canopy itself, but at and beneath the forest floor as well.     

Nitrogen fixation associated with the cyanobacterial mat of a marine laminated microbial ecosystem

The nitrogenase activity in the cyanobacterial mat of a laminated microbial ecosystem was investigated by the acetylene reduction method. Measurements under several conditions such as light and dark, aerobic and anaerobic and by inhibiting photosystem II by 10^-5 M DCMU showed the nitrogenase activity to be light stimulated and to some degree inhibited by oxygen. An appreciable amount of activity was also present under complete aerobic conditions. We estimated 8 to 15 kg N fixed per hectare per year for that part of the intertidal flat supporting growth of cyanobacteria. By measuring a vertical sediment profile, nitrogenase was shown to be associated with the cyanobacterial mat. Diurnal measurements of nitrogenase showed two activity peaks, one at sunrise and one at sunset. Following population dynamics in the cyanobacterial mat showed Microcoleus sp., Oscillatoria spp., Spirulina sp., Gloeocapsa sp. and sometimes Merismopedia sp. to be present. During four years of observations we never found any heterocystous cyanobacteria. Non-heterocystous cyanobacteria apparently play an important role in nitrogen fixation in this marine intertidal environment.     

Evidence of nitrification and denitrification in high and low microbial abundance sponges

Aerobic and anaerobic microbial key processes were quantified and compared to microbial numbers and morphological structure in Mediterranean sponges. Direct counts on histological sections stained with DAPI showed that sponges with high microbial abundances (HMA sponges) have a denser morphological structure with a reduced aquiferous system compared to low microbial abundance (LMA) sponges. In Dysidea avara, the LMA sponge, rates of nitrification and denitrification were higher than in the HMA sponge Chondrosia reniformis, while anaerobic ammonium oxidation and sulfate reduction were below detection in both species. This study shows that LMA sponges may host physiologically similar microbes with comparable or even higher metabolic rates than HMA sponges, and that anaerobic processes such as denitrification can be found both in HMA and LMA sponges. A higher concentration of microorganisms in the mesohyl of HMA compared to LMA sponges may indicate a stronger retention of and, hence, a possible benefit from associated microbes.     

N fertilization effects on denitrification and N cycling in an aggrading forest.

We investigated N cycling and denitrification rates following five years of N and dolomite amendments to whole-tree harvested forest plots at the long-term soil productivity experiment in the Fernow Experimental Forest in West Virginia, USA. We hypothesized that changes in soil chemistry and nutrient cycling induced by N fertilization would increase denitrification rates and the N2O:N2 ratio. Soils from the fertilized plots had a lower pH (2.96) than control plots (3.22) and plots that received fertilizer and dolomite (3.41). There were no significant differences in soil %C or %N between treatments. Chloroform-labile microbial biomass carbon was lower in fertilized plots compared to control plots, though this trend was not significant. Extractable soil NO3- was elevated in fertilized plots on each sample date. Soil-extractable NH4+, NO3-, pH, microbial biomass carbon, and %C varied significantly by sample date suggesting important seasonal patterns in soil chemistry and N cycling. In particular, the steep decline in extractable NH4+ during the growing season is consistent with the high N demands of a regenerating forest. Net N mineralization and nitrification also varied by date but were not affected by the fertilization and dolomite treatments. In a laboratory experiment, denitrification was stimulated by NO3- additions in soils collected from all field plots, but this effect was stronger in soils from the unfertilized control plots, suggesting that chronic N fertilization has partially alleviated a NO3- limitation on denitrification rates. Dextrose stimulated denitrification only in the whole-tree-harvest soils. Denitrification enzyme activity varied by sample date and was elevated in fertilized plots for soil collected in July 2000 and June 2001. There were no detectable treatment effects on N2O or N2 flux from soils under anaerobic conditions, though there was strong temporal variation. These results suggest that whole-tree harvesting has altered the N status of these soils so they are less prone to N saturation than more mature forests. It is likely that N losses associated with the initial harvest and high N demand by aggrading vegetation is minimizing, at least temporarily, the amount of inorganic N available for nitrification and denitrification, even in the fertilized plots in this experiment.     

The role of microbial mats during primary succession in calcareous dune slacks: an experimental approach

Laminated microbial mats from a sandy beach plain were grown in water-saturated pots in a glass house for six months and then used to assess their effect on the establishment of juveniles of three plant species representing different successional stages in dune slack development. The selected species wereSamolus valerandi, characteristic of pioneer stages,Calamagrostis epigejos, characteristic of more productive, late successional stages, andJuncus alpinoarticulatus, which occurs in a wide range of successional stages. Juveniles of all three species that were placed on top of intact living microbial mats established themselves in the mat.C. epigejos andJ. alpinoarticulatus survived for several weeks but later on their numbers decreased and the total biomass production of the species after six months was poor.S. valerandi, in contrast, grew profusely in intact microbial and algal mats. Heating of the microbial mat by heat sterilization, prior to the experiment, did not improve the performance of the species. When the juveniles were planted in the microbial mats after breaking the surface of the mat, the survival of juveniles ofC. epigejos andJ. alpinoarticulatus was much higher and so was the biomass of surviving plants after six months. Planting ofSamolus in the mats had some positive effect on the survival percentage of the juveniles, but not on the total biomass at the end of the experiment. Slightly lower water tables had a negative effect on the performance of all species. Measurements of the pH in the pots revealed that there were no significant differences in the top layer. Sulphide concentrations were very low in all the pots where juveniles had been planted and also in the pots withS. valerandi. Relatively high concentrations (30–50 μmol/l) were found in pots with poor growth ofJuncus andCalamagrostis plants. These values may exceed toxic levels for these species. Although oxygen concentrations in the pots were generally low, no relation existed between plant biomass and oxygen content, indicating that plant growth was not primarily limited by oxygen stress. These experiments support the idea that microbial mats may assist in extending the life span of early pioneer stages during dune slack succession by inhibiting the growth of species of later successional stages.     

Seasonal variations in nitrogen-fixation (acetylene reduction) and sulphate-reduction rates in the rhizosphere of Zostera noltii: nitrogen fixation by sulphate-reducing bacteria

Nitrogen-fixation (acetylene reduction) rates were measured over an annual cycle in meadows of the seagrass Zostera noltii Hornem in the Bassin d'Arcachon, south-west France, between March 1994 and February 1995, using both slurry and whole-core techniques. Measured rates using the slurry technique consistently overestimated those determined on whole cores, probably due to the release of labile organic carbon sources as a result of root damage during preparation of the slurries. Thus, the whole-core technique may provide a more accurate estimate of in situ activity, since disturbance of physicochemical gradients of oxygen, sulphide, nutrients and the relationship between the plant roots and the rhizosphere microflora is minimised. Rates measured by the whole-core method were 1.8- to 4-fold greater (dependent upon season) in the light than those measured during dark incubations, indicating that organic carbon diffusing from the plant roots during photosynthesis was an important factor in regulating nitrogen fixation in the rhizosphere. Additions of sodium molybdate, a specific inhibitor of sulphate-reducing bacteria (SRB) inhibited acetylene-reduction activity by >80% as measured by both the slurry and whole-core techniques throughout the year, inferring that SRB were the dominant component of the nitrogen-fixing microflora. A mutualistic relationship between Z. noltii and nitrogen-fixing SRB in the rhizosphere, based on the exchange of organic carbon and fixed nitrogen is proposed. Acetylene- and sulphate-reduction rates showed distinct summer peaks which correlated with a reduced availability of ammonium in the sediment and the annual growth cycle of Z. noltii in the basin. Overall, these data indicate that acetylene reduction (nitrogen fixation) activity in the rhizosphere of Z. noltii was regulated both by the availability of organic carbon from the plant roots and maintenance of a low NH ₄ ⁺ concentration in the vicinity of the plant roots due to efficient assimilation of NH ₄ ⁺ by Z. noltii during the growth season. Nitrogenfixation rates determined from acetylene-reduction rates measured using the whole-core technique ranged from 0.1 to 7.3 mg N m^-2d^-1, depending on season, and were calculated to contribute between 0.4 and 1.1 g N m^-2yr^-1, or 6.3 to 12% of the annual fixed nitrogen requirement of Z. noltii.     

Floodplain restoration enhances denitrification and reach-scale nitrogen removal in an agricultural stream

Streams of the agricultural Midwest, USA, export large quantities of nitrogen, which impairs downstream water quality, most notably in the Gulf of Mexico. The two-stage ditch is a novel restoration practice, in which floodplains are constructed alongside channelized ditches. During high flows, water flows across the floodplains, increasing benthic surface area and stream water residence time, as well as the potential for nitrogen removal via denitrification. To determine two-stage ditch nitrogen removal efficacy, we measured denitrification rates in the channel and on the floodplains of a two-stage ditch in north-central Indiana for one year before and two years after restoration. We found that instream rates were similar before and after the restoration, and they were influenced by surface water NO3- concentration and sediment organic matter content. Denitrification rates were lower on the constructed floodplains and were predicted by soil exchangeable NO3- concentration. Using storm flow simulations, we found that two-stage ditch restoration contributed significantly to NO3- removal during storm events, but because of the high NO3- loads at our study site, < 10% of the NO3- load was removed under all storm flow scenarios. The highest percentage of NO3- removal occurred at the lowest loads; therefore, the two-stage ditch's effectiveness at reducing downstream N loading will be maximized when the practice is coupled with efforts to reduce N inputs from adjacent fields.     

Nitrogen fixation in the North Pacific Ocean

Nitrogen fixation in the euphotic zone of the ocean was measured by C₂H₂ reduction and ¹⁵N₂ incorporation associated with Trichodesmium sp. and also with Richelia intracellularis occurring within the cells of Rhizosolenia styliformis var. longispina, and R. cylindrus. The vertical distribution of N₂ fixation activity, N₂-fixing species, particulate matter and dissolved nutrients was measured. The effects of light intensity, sample concentration, length of incubation, and nutrient enrichment on the rates of C₂H₂ reduction were determined. Estimates of the importance of N₂ fixation in adding previously uncycled nitrogen to the euphotic zone are given.     

Nutrient cycling studies in Carmarthen Bay: phytoplankton production,nitrogen assimilation and regeneration

The results from a series of five cruises (August 1979–January 1981) to examine rates of primary production, nitrogen assimilation and regeneration in Carmarthen Bay, S. Wales, are presented. Chlorophyll, primary production and regeneration were distributed irregularly throughout the bay, with the highest concentrations and rates being found around the shallow periphery. Nitrate was the dominant form of nitrogen assimilated by the phytoplankton, apart from periods of high biomass during the summer when ammonium assumed a more important role. The rates of ammonium regeneration from microheterotrophs were greater than the ammonium demand over much of the year, indicating that the bay was a net exporter of ammonium and was a physically dominated open system. During the summer, the rates of ammonium regeneration and assimilation were similar, indicating efficient recycling of nutrients.     

Nitrogen fixation in the rhizosphere of marine angiosperms

High rates of acetylene reduction were observed in systems containing excised rhizomes of the Caribbean marine angiosperms Thalassia testudinum, Syringodium filiforme and Diplanthera wrightii, and the temperate marine angiosperm Zostera marina. For 4 plant and plant-sediment systems the ratio of acetylene reduced/N₂ fixed varied from 2.6 to 4.6. For T. testudinum the estimated rates of nitrogen fixation are in agreement with estimated requirements of the plant for nitrogen. For a typical T. testudinum stand, N₂ fixation is estimated to be 100 to 500 kg N/hectare per year. Numbers of N₂-fixing bacteria in the rhizosphere sediments were roughly 50 to 300 times more abundant than those in the nonrhizosphere sediments, and in both types of sediments were of the same orders as the estimated numbers of heterotrophic aerobes.Canadian IBP Contribution No. 137.     

Nitrogen cycling in a marine planktonic food chain: Nitrogen fluxes through the principal components and the effects of adding copper

Nitrogen cycling in a natural planktonic food chain was followed before and after perturbation by copper (10 g l^-1). Observations were separated into effects from artificial containment of the food chain and effects from the addition of copper. An initial reduction of seston-N and subsequent recovery, uniformly high settling rates of suspended particulate-N, and general decrease in zooplankton-N were attributed to the physical limitations imposed by the enclosures (e.g. elimination of normal mixing processes). Acute inhibition of nitrate assimilation by the seston, acute elevation of seston-N settling and chronic inhibition of zooplankton feeding, along with accelerated decline in zooplankton stocks, were attributed to copper stress. Since zooplankton stocks and fluxes were most drastically affected by copper, its overall effect on N-cycling through the planktonic food chain would depend largely upon the importance of zooplankton in regulating the production of the remaining seston.     

Stoichiometric controls of nitrogen and phosphorus cycling in decomposing beech leaf litter

Resource stoichiometry (C:N:P) is an important determinant of litter decomposition. However, the effect of elemental stoichiometry on the gross rates of microbial N and P cycling processes during litter decomposition is unknown. In a mesocosm experiment, beech (Fagus sylvatica L.) litter with natural differences in elemental stoichiometry (C:N:P) was incubated under constant environmental conditions. After three and six months, we measured various aspects of nitrogen and phosphorus cycling. We found that gross protein depolymerization, N mineralization (ammonification), and nitrification rates were negatively related to litter C:N. Rates of P mineralization were negatively correlated with litter C:P. The negative correlations with litter C:N were stronger for inorganic N cycling processes than for gross protein depolymerization, indicating that the effect of resource stoichiometry on intracellular processes was stronger than on processes catalyzed by extracellular enzymes. Consistent with this, extracellular protein depolymerization was mainly limited by substrate availability and less so by the amount of protease. Strong positive correlations between the interconnected N and P pools and the respective production and consumption processes pointed to feed-forward control of microbial litter N and P cycling. A negative relationship between litter C:N and phosphatase activity (and between litter C:P and protease activity) demonstrated that microbes tended to allocate carbon and nutrients in ample supply into the production of extracellular enzymes to mine for the nutrient that is more limiting. Overall, the study demonstrated a strong effect of litter stoichiometry (C:N:P) on gross processes of microbial N and P cycling in decomposing litter; mineralization of N and P were tightly coupled to assist in maintaining cellular homeostasis of litter microbial communities.     

氮输入对纳帕海沼泽湿地土壤氨挥发和反硝化的影响

选取纳帕海常见湿地植物茭草（Zizania caduciflora）和水葱（Scirpus validus）]及其生长土壤为对象,通过培养实验,研究了3个不同氮输入水平[0g-m^2（对照,NO）、20g-m^2（N20）、40g-m^2。（N40）]对茭草和水葱湿地土壤氨挥发和反硝化的影响。结果表明：氮输入促进了茭草和水葱湿地土壤氨挥发化作用,增加了湿地土壤氨挥发的累积量。适量的氮输入对湿地土壤氨挥发促进显著。在培养前期适量氮输入下的氨挥发积累量高于高水平氮输入的,随着培养时间的延长,后期适量氮输入下的积累量明显下降;而高水半氮输入处理下氨挥发积累量的明显增加,适量氮输入下茭草氨挥发积累大于水葱。氮输入增强了菱草和水葱湿地土壤反硝化作用,加快了反硝化N2O的排放。适量的氮输入促进茭草反硝化作用和反硝化损失量增加明显,培养的前期的适量氮输入处理下的反硝化作用和反硝化N2O的排放增强不明显,随着培养时间的延长,高水平氮输入处理下的反硝化损失量越明显,并且水葱较茭草更为明显。后期适量氮输入下的反硝化损失速率和反硝化损失量高于高水平氮输入,适量氮输入较高水平氮输人促进明显,高水平的氮输入限制了反硝化损失,反硝化N2O的排放总量下降,土壤中氮富集增大。     

西藏高原农业生态系统氮素循环的数量特征研究

以西藏贡嘎县为研究对象，运用生态学原理，定量估算了西藏高原农业生态系统中氮素循环的数量特征，为该系统的调控与结构优化提供了理论依据。     

Nitrogen and phosphorus cycling in shrimp ponds and the measures for sustainable management

Six ponds of age 3 were selected 45 km north from Suzhou in the Tailake region, and research conducted on nitrogen and phosphorus cycling in P. vannanmei (Penaeus vannanme) ponds and M. nipponense (Macrobrachium nipponense) hatchery ponds under normal management. Two treatments each had three replications. The results confirmed that feed was the major path of nitrogen and phosphorus input, each accounted for 61.24% (193.81 kg ha(-1)) and 81.08% (45.20 kg ha(-1)) of the total nitrogen and phosphorus input for P. vannanme ponds; the values for M. nipponense ponds were 43.93% (86.31 kg ha(-1)) and 57.67% (14.61 kg ha(-1)), respectively. Water pumped into ponds contributed on average 83.57 kg ha(-1) nitrogen and 8.48 kg ha(-1) phosphorus for P. vannanmei ponds, and 87.48 kg ha(-1) nitrogen and 7.00 kg ha(-1) phosphorus for M. nipponense hatchery ponds. Shrimp harvest recovered 102.81 kg ha(-1) nitrogen (32.94% of the total nitrogen input) and 7.94 kg ha(-1) phosphorus (14.23% of the total phosphorus input) for P. vannanme ponds; and 43.94 kg ha(-1) nitrogen and 4.46 kg ha(-1) phosphorus for M. nipponense hatchery ponds. The sum of nitrogen losses through volatilization, denitrification and sedimentation was 173.62 and 122.39 kg ha(-1), 54.86% and 62.29% of the total nitrogen input for P. vannanme ponds and M. nipponense hatchery ponds, respectively. Sediment accumulated 41.46 and 14.63 kg ha(-1) phosphorus, 74.37% and 64.85% of the total phosphorus input for P. vannanm ponds and M. nipponense hatchery ponds. Draining and seeping caused 40.06 kg ha(-1) nitrogen (12.66% of total nitrogen input) and 6.36 kg ha(-1) phosphorus (11.40% of total phosphorus input) loss to the surrounding water from P. vannanme ponds in 114 days; 30.14 kg ha(-1) nitrogen (15.34% of the total input) and 4.45 kg ha(-1) phosphorus (17.57% of the total input) to channel water from M. nipponense hatchery ponds in 87 days, respectively. Countermeasures for sustainable pond management include improving feeds and feeding, sediment treatments, machine aerating, chemicals with no pollution, and integrated fish-shrimp cultivation. Management of water resources for pond and methods to reduce nitrogen and phosphorus loading into surrounding water from drainage are elucidated.     

Desiccation enhanced nitrogen uptake rates in intertidal seaweeds

Desiccation increased nitrate and ammonium uptake rates upon resubmergence in late summer populations of the intertidal macroalgae Gigartina papillata (C.Ag.) J.Ag., Enteromorpha intestinalis (L.) Grev., Fucus distichus L., and Pelvetiopsis limitata (Setch) Gardn. The ratio of nitrogen uptake rates in desiccated plants to rates in hydrated plants (controls) was correlated with the position of the species in the intertidal zone. Gracilaria pacifica Abbott., the species occurring at the lowest shore level, showed no enhancement of nitrogen uptake following desiccation. The high intertidal species such as P. limitata and F. distichus showed a two-fold enhancement of nitrate and ammonium uptake following more extensive desiccation (>30%) and continued uptake even following severe desiccation (50 to 60%). After the plants had been desiccated, the increase in nitrate uptake rates upon submergence lasted much longer than a similar enhancement of ammonium uptake. The duration of the enhanced nitrate uptake was similar to the time required for total rehydration but the uptake rates were not related to the state of rehydration. The potential contribution that this enhanced nitrogen uptake following desiccation could make to total nitrogen procurement for growth is discussed. The experiments were carried out in 1979 or 1980 and repeated in 1981.This paper is dedicated to Dr. R. F. Scagel on the occasion of his retirement for his outstanding contribution to phycology     

Carbon and nitrogen metabolism by Monochrysis lutheri: Measurement of growth-rate-dependent respiration rates

Dark respiration rates were measured and carbon-excretion rates calculated for a nitrate-limited population of the marine chrysophyte Monochrysis lutheri grown in continuous culture at 20°C on a 12 h light-12 h dark cycle of illumination and over a series of 4 growth rates. A significant (P<0.05) positive correlation was found between dark respiration rate and growth rate. From a simple linear fit to the data, the respiration rate at maximum growth rate was estimated to be roughly 10.5% of the maximum gross-carbon-production rate, and more than three times higher than the extrapolated respiration rate at zero net-growth rate. Carbon-excretion rates showed no significant correlation with growth rate, and averaged less than 5% of the maximum gross-carbon-production rate. Mean cell nitrogen to carbon ratios were correlated in a virtually linear manner (r=0.994) with growth rate, and at a given growth rate were consistently higher than nitrogen to carbon ratios for the same species grown on continuous light. A comparison of carbon and nitrogen quotas as a function of growth rate for M. lutheri and other species suggests that the increase of cellular nitrogen at high growth rates under nitrate-limited growth conditions may be associated with the storage of cellular protein or amino acids rather than the presence of an inorganic nitrogen reservoir. The maximum nitrate uptake rate per cell during the day changed very little over the range of growth rates studied, and was comparable to the maximum uptake rate found for cells grown on continuous light. However, the cell nitrogen quota increased steadily with growth rate, causing a reduction in the maximum specific-uptake rate of nitrate during the day at high growth rates. The dark nitrate-uptake capacity of the population was clearly exceeded by the supply rate at the two higher growth rates, leading to a buildup of nitrate during the night which amounted to as much as 21% of the particulate nitrogen in the growth chamber by morning.Hawaii Institute of Marine Biology Contribution No. 478.     

灰泥土中不同氮肥品种反硝化损失与N2O排放量的差异

在实验室培养条件下，采用土壤培养-乙炔抑制法测定尿素、碳酸氢铵、硫酸铵和氯化铵4种氮肥品种在灰泥土中反硝化损失和N2O排放量的差异。结果表明，氮肥品种间的N2O排放量和反硝化损失量存在极显著差异。尿素、碳铵和氯化铵的N2O排放量极显著高于硫铵，占施肥量的3．88%-4．14％；尿素和碳铵的反硝化损失量分别为施肥量的5．8％和3．7％，极显著高于硫铵和氯化铵；但氯化铵的反硝化损失量显著低于CK。     

Resistance of nutrient cycling systems to perturbations of the flow rates

The resistance of a nutrient cycling system to a disturbance that alters the flow rates, measured by the displacement of the nutrient levels while the disturbance continues, is as important an aspect of stability as the rate of return to equilibrium after the disturbance ends. Two important characteristics of the resistance of a system are the ultimate displacement that would occur if the disturbance lasted indefinitely, and the rate at which the system approaches the ultimate displacement.The mathematical theory is worked out for linear systems and applied to a group of eight nutrient cycling models. The relative resistance of individual compartments of different systems depends heavily on the period of interest, because of the wide range of magnitudes of the eigenvalues, which determine the response times. However, when measuring the change in total nutrients in a system with small exterior flows compared with the amount of internal cycling, only the critical root is significant in determining the response rate.     

岩溶生态系统中土壤微生物量氮含量及其变化规律

选择果树园和林地两种土地利用类型来研究土壤微生物量氮(Soil microbial biomass nitrogen,SMBN)含量及其与土壤溶解有机氮(Soil dissolved organic nitrogen,DON)、碱解氮的关系。结果表明:SMBN含量范围在5.33~34.61kg·hm-2之间,平均为14.81kg·hm-2,仅为文献中报道的SMBN的3%~37%。SMBN占土壤全氮的比例较小,为0.25%~1.5%,平均为0.754%。SMBN与DON及土壤有机碳等指标变化趋势一致。SMBN易矿化,与碱解氮关系密切,但由于岩溶山区这种特殊的生态环境,使得这种关系不像文献中报道的呈极强的正相关关系,这对于研究岩溶区特定条件下SMBN的变化以及对当地的土地利用类型有一定的参考价值和现实意义。