INTERACTION OF NITROGEN FIXATION AND PHOSPHORUS LIMITATION IN APHANIZOMENON FLOS-AQUAE (CYANOPHYCEAE)1

The effect of simultaneous nitrogen fixation and phosphorus limitation on the physiological adaptation and growth performance of Aphanizomenon flos-aquae (L.) Ralfs PCC 7905 was studied in continuous culture. In the absence of ammonia, N₂ fixation occurred and the maximum growth rate (as determined in diluted batch cultures) was lower. However, no distinction could be made between the steady-state N uptake rates (based on cellular N contents) of N₂-fixing cells and cells grown with ammonia. At the higher dilution rates, the residual P concentration increased with increasing dilution rate, more so under N₂-fixing conditions, compared to the cultures grown in the presence of ammonia. More generally, the yield of biomass per consumed P, as the biomass concentration itself, decreased with increasing dilution rate, and both were lower under N₂-fixing conditions. The restricted biomass production under N₂-fixing conditions suggests that reduction of N loading may benefit lake restoration projects. The influence of N₂-fixation on the severity of P limitation is discussed in terms of metabolic control analysis. From the increase of the residual P concentration on switching from ammonium to N₂-fixing conditions, it is deduced that under N₂-fixing and P-limited conditions, control of growth is shared by N and P metabolism.     

The impact of simulated chronic nitrogen deposition on the biomass and N2-fixation activity of two boreal feather moss–cyanobacteria associations

Bryophytes achieve substantial biomass and play several key functional roles in boreal forests that can influence how carbon (C) and nitrogen (N) cycling respond to atmospheric deposition of reactive nitrogen (N_r). They associate with cyanobacteria that fix atmospheric N₂, and downregulation of this process may offset anthropogenic N_r inputs to boreal systems. Bryophytes also promote soil C accumulation by thermally insulating soils, and changes in their biomass influence soil C dynamics. Using a unique large-scale (0.1 ha forested plots), long-term experiment (16 years) in northern Sweden where we simulated anthropogenic N_r deposition, we measured the biomass and N₂-fixation response of two bryophyte species, the feather mosses Hylocomium splendens and Pleurozium schreberi. Our data show that the biomass declined for both species; however, N₂-fixation rates per unit mass and per unit area declined only for H. splendens. The low and high treatments resulted in a 29% and 54% reduction in total feather moss biomass, and a 58% and 97% reduction in total N₂-fixation rate per unit area, respectively. These results help to quantify the sensitivity of feather moss biomass and N₂ fixation to chronic N_r deposition, which is relevant for modelling ecosystem C and N balances in boreal ecosystems.     

Effect of substrate nitrogen on the performance ofin vitro propagatedAlnus glutinosa clones inoculated with Sp+ and Sp− Frankia strains

The addition of combined nitrogen to substrate at an appropriate rate can stimulate N₂-fixation thus inreasing the efficiency of the Alnus-Frankia symbiosis. To examine how nitrogen additions can effect the peformance of different pairs of symbionts, growth and time course of N₂-fixation were studied in plants supplied with NH₄NO₃. Two cloned ofAlnus glutinosa (L.) Gaertn., propagatedin vitro, were inoculated with two strains ofFrankia (AVP3d and ACN14a) and grown in a greenhouse. Calcined montmorillonite (Totfaice^R) was used as growth substrate. Six N treatments were made up of varied amounts of NH₄NO₃ supplied in one single addition shortly before inoculation. Weekly measurements of shoot height and repeated measurements of nitrogenase activity (acetylene reduction) performed on intact root systems were used to monitor the development of the symbioses. Nitrogen treatments containing from 0.10 to 0.68 mg N g⁻¹ dry substrate stimulated N₂-fixation as well as growth. The relative performance of the two clones was different according to N treatment; one clone showed a greater benefit from the nitrogen input. Our results support the recommendation that selection of symbionts according to performance should be carried out with an input of combined nitrogen. This can provide optimum conditions for the development of each pair of symbionts.     

Utilization of carbon and nitrogen reserves of alfalfa roots in supporting N2-fixation and shoot regrowth

Perennial legume such as alfalfa have the capacity to sustain shoot regrowth and some nodule N₂-fixation after removal (cutting) of shoots which contain practically all of the plant's photosynthetic capacity. The role of the roots in supporting these processes has not been fully described. Measurements were made of the nodules' responses to removal of shoots from 8-week-old seedlings in terms of N₂-fixation, as nitrogenase activity (NA) measured as acetylene reduction, dark CO₂ fixation, measured as in vitro phosphoenolpyruvate carboxylase (PEPC) activity, and total non-structural carbohydrate (NSC) content. These properties decreased and recovered in that sequence, which suggests that nodule NSC supported the substrate requirements of NA and PEPC immediately after cutting. The utilization and redistribution or root carbon and nitrogen, prelabeled with ¹⁴C and ¹⁵N, were also followed after cutting 8-week-old alfalfa seedlings. In the first 2 weeks of regrowth 12% of root C and 25% of root N were transferred for incorporation into new shoots. Up to 40% of the root C was used for plant respiration to support 28 days of shoot regrowth and N₂-fixation. The decline of N₂-fixation was slower after cutting and its minimum activity rose up 45% of pre-cut activity as root reserves were built up with plant age. Therefore, the stored reserves of nodulated roots play an important role in support of N₂-fixation after cutting.Contribution No. 1265 from Plant Research Center.Contribution No. 1265 from Plant Research Center.     

Nitrogen dynamics in two antarctic streams

The many glacier meltwater streams of southern Victoria Land flow through catchments where life forms are almost entirely microbial. Allochthonous inputs of nitrogen from two study streams near McMurdo Sound were derived mostly from the melting glaciers (ca. 100–200 mg N m^–3) with some originating from N₂-fixation by heterocystous cyanobacteria (max. 939 mg N m^–2 year^–1). Thirty to fifty per cent of the glacier derived N was dissolved organic N and a major proportion of this was identified as urea N which was utilised by the rich algal and cyanobacterial mats in the streams. A nutrient budget for Fryxell Stream was estimated, quantifying uptake of urea-N and dissolved inorganic N and the release of dissolved organic (non urea) and particulate N by the stream communities. An index of in-stream nitrogen processing, the Net Uptake Length Constant in these streams was compared with that from temperate climates and was found to be similar. Despite the influence of low temperatures on microbial activity (mean daily water temperature = 5 °C) nutrient removal rates from these antarctic streams are high because of the large standing stock of microbial biomass there.     

Short-term Nitrogen Fixation by Legume Seedlings and Resprouts After Fire in Mediterranean Old-fields

Fires may greatly alter the N budget of a plant community. During fire nitrogen is lost to the atmosphere. Although high light availability after fire promotes N₂-fixation, the presumably high soil N availability could limit N₂-fixation activity. The latter limitation might be particularly acute in legume seedlings compared with resprouts, which have immediate access to belowground stored carbon. We wished to learn whether early post-fire conditions were conducive to N₂-fixation in leguminous seedlings and resprouts in two types of grassland and in a shrubland and whether seedlings and resprouts incurred different amounts of N₂-fixation after fire. We set 18 experimental fires in early autumn on 6 plots, subsequently labelling 6 subplots (2 × 2 m²) in each community with ¹⁵NH₄⁺-N (99 atom % excess). For 9 post-fire months we measured net N mineralisation in the top 5 cm of soil and we calculated the fraction of legume N derived from the atmosphere (%Ndfa) in seedlings and resprouts. We used two independent estimates of the amounts of N derived from non-atmospheric sources in potentially N₂-fixing plants: mean soil pool abundance and the ¹⁵N-enrichment of non-legumes. Despite substantial soil net N mineralisation in all burned community types (about 2.6 g Nm⁻² during the first nine months after fire), the %Ndfa of various legume species was 52–99%. Legumes from both grasslands showed slightly higher N₂-fixation values than shrubland legumes. As grassland legumes grew in more belowground dense communities than shrubland legumes, we suggest that higher competition for soil resources in well established grass-resprouting communities may enhance the rate of N₂-fixation after fire. In contrast to our hypothesis, legume seedlings and resprouts from the three plant communities studied, had similar %Ndfa and apparently acquired most of their N from the atmosphere rather than from the soil.     

Isotopic evidence for increased carbon and nitrogen exchanges between peatland plants and their symbiotic microbes with rising atmospheric CO2 concentrations since 15,000 cal. year BP

Whether nitrogen (N) availability will limit plant growth and removal of atmospheric CO₂ by the terrestrial biosphere this century is controversial. Studies have suggested that N could progressively limit plant growth, as trees and soils accumulate N in slowly cycling biomass pools in response to increases in carbon sequestration. However, a question remains over whether longer-term (decadal to century) feedbacks between climate, CO₂ and plant N uptake could emerge to reduce ecosystem-level N limitations. The symbioses between plants and microbes can help plants to acquire N from the soil or from the atmosphere via biological N₂ fixation—the pathway through which N can be rapidly brought into ecosystems and thereby partially or completely alleviate N limitation on plant productivity. Here we present measurements of plant N isotope composition (δ¹⁵N) in a peat core that dates to 15,000 cal. year BP to ascertain ecosystem-level N cycling responses to rising atmospheric CO₂ concentrations. We find that pre-industrial increases in global atmospheric CO₂ concentrations corresponded with a decrease in the δ¹⁵N of both Sphagnum moss and Ericaceae when constrained for climatic factors. A modern experiment demonstrates that the δ¹⁵N of Sphagnum decreases with increasing N₂-fixation rates. These findings suggest that plant-microbe symbioses that facilitate N acquisition are, over the long term, enhanced under rising atmospheric CO₂ concentrations, highlighting an ecosystem-level feedback mechanism whereby N constraints on terrestrial carbon storage can be overcome.     

Glucose Metabolism in Batch and Continuous Cultures of Gluconacetobacter diazotrophicus PAL 3

Periplasmic glucose oxidation (by way of a pyrrolo-quinoline-quinone [PQQ]–linked glucose dehydrogenase [GDH]) was observed in continuous cultures of Gluconacetobacter diazotrophicus regardless of the carbon source (glucose or gluconate) and the nitrogen source (N₂ or NH₃). Its synthesis was stimulated by conditions of high energetic demand (i.e., N₂-fixation) and/or C-limitation. Under C-excess conditions, PQQ-GDH synthesis increased with the glucose concentration in the culture medium. In batch cultures, PQQ-GDH was actively expressed in very early stages with higher activities under conditions of N₂-fixation. Hexokinase activity was almost absent under any culture condition. Cytoplasmic nicotinamide adenine dinucleotide (NAD)–linked glucose dehydrogenase (GDH) was expressed in continuous cultures under all tested conditions, and its synthesis increased with the glucose concentration. In contrast, low activities of this enzyme were detected in batch cultures. Periplasmic oxidation, by way of PQQ-GDH, seems to be the principal pathway for metabolism of glucose in G. Diazotrophicus, and NAD-GDH is an alternative route under certain environmental conditions.     

Heterotrophic nitrogen removal by <Emphasis Type="Italic">Providencia rettgeri</Emphasis> strain YL

Providencia rettgeri strain YL was found to be efficient in heterotrophic nitrogen removal under aerobic conditions. Maximum removal of NH₄ ⁺–N occurred under the conditions of pH 7 and supplemented with glucose as the carbon source. Inorganic ions such as Mg²⁺, Mn²⁺, and Zn²⁺ largely influenced the growth and nitrogen removal efficiency. A quantitative detection of nitrogen gas by gas chromatography was conducted to evaluate the nitrogen removal by strain YL. From the nitrogen balance during heterotrophic growth with 180 mg/l of NH₄ ⁺–N, 44.5% of NH₄ ⁺–N was in the form of N₂ and 49.7% was found in biomass, with only a trace amount of either nitrite or nitrate. The utilization of nitrite and nitrate during the ammonium removal process demonstrated that the nitrogen removal pathway by strain YL was heterotrophic nitrification-aerobic denitrification. A further enzyme assay of nitrate reductase and nitrite reductase activity under the aerobic condition confirmed this nitrogen removal pathway.     

Bryophyte-cyanobacterial associations as a key factor in N2-fixation across the Canadian Arctic

Nitrogen inputs via biological N₂-fixation are important in arctic environments where N often limits plant productivity. An understanding of the direct and indirect theoretical causal relationships between key intercorrelated variables that drive the process of N₂-fixation is essential to understanding N input. An exploratory multi-group Structural Equation Modeling (SEM) approach was used to examine the direct and indirect effects of soil moisture, plant community functional composition, and bryophyte and lichen abundance on rates of nitrogen fixation at a low arctic ecosystem, two high arctic oases and a high arctic polar desert in the Canadian Arctic. Increasing soil moisture was strongly associated with an increasing presence of bryophytes and increasing bryophyte abundance was a major factor determining higher N₂-fixation rates at all sites. Shrubs had a negative effect on bryophyte abundance at all sites with the exception of the polar desert site at Alexandra Fjord highland. The importance of competition from vascular plants appears to be greater in more productive sites and may increase at lower latitudes. Moisture availability may have an indirect effect on ecosystem development by affecting N input into the system with bryophyte-cyanobacterial associations playing an important intermediary role in the process.     

Growth and nitrogen acquisition strategies of Acacia senegal seedlings under exponential phosphorus additions

There remains conflicting evidence on the relationship between P supply and biological N₂-fixation rates, particularly N₂-fixing plant adaptive strategies under P limitation. This is important, as edaphic conditions inherent to many economically and ecologically important semi-arid leguminous tree species, such as Acacia senegal, are P deficient. Our research objective was to verify N acquisition strategies under phosphorus limitations using isotopic techniques. Acacia senegal var. senegal was cultivated in sand culture with three levels of exponentially supplied phosphorus [low (200 μmol of P seedling⁻¹ over 12 weeks), mid (400 μmol) and high (600 μmol)] to achieve steady-state nutrition over the growth period. Uniform additions of N were also supplied. Plant growth and nutrition were evaluated. Seedlings exhibited significantly greater total biomass under high P supply compared to low P supply. Both P and N content significantly increased with increasing P supply. Similarly, N derived from solution increased with elevated P availability. However, both the number of nodules and the N derived from atmosphere, determined by the ¹⁵N natural abundance method, did not increase along the P gradient. Phosphorus stimulated growth and increased mineral N uptake from solution without affecting the amount of N derived from the atmosphere. We conclude that, under non-limiting N conditions, A. senegal N acquisition strategies change with P supply, with less reliance on N₂-fixation when the rhizosphere achieves a sufficient N uptake zone.     

Nitrogen fixation varies spatially and seasonally in linked stream-lake ecosystems

We performed surveys of nitrogen (N₂)-fixation in three oligotrophic lake-stream systems in the Sawtooth Mountains of central Idaho to address two questions: (1) Which habitat types within linked lake-stream systems (lake pelagic, lake benthic, and stream) exhibit the highest rates of N₂ fixation?, and (2) How does N₂ fixation compare to the hydrologic flux of nitrogen? A seasonal survey showed that N₂ fixation in a single lake and its outlet stream peaked in late summer, when hydrologic N fluxes were lowest. Benthic lake N₂-fixation rates by epiphytes were highest at mid-lake depths, where their percent cover was highest, while rates by epipelon were greatest at shallow lake depths. Pelagic N₂ fixation was below detection. Stream N₂-fixation rates were greatest on rock substrates and in the lake outlet stream. These patterns were supported by a baseflow survey (late July) in three lake-stream ecosystems which confirmed that N₂-fixation rates peaked in the lake benthos at shallow depths and on rock substrates in outlet streams. Scaling N₂-fixation rates to whole lake and stream areas revealed that N₂ fixation could exceed the nitrate, and sometimes the total dissolved nitrogen flux during baseflow in lakes and outlet streams. Despite low rates, total N₂-fixation contributions (kg/day) from lakes were greater because they had far larger surface areas than the stream environments. Fixed nitrogen contributions from stream outlets were also relatively high because of high N₂-fixation rates and despite low surface areas. This study suggests that N₂ fixation could be a seasonally important nitrogen source to nutrient deficient subalpine lake-stream ecosystems. In addition, the frequency and location of lakes could control N₂-fixation contributions to watersheds by providing a large area for within-lake N₂ fixation, and creating conditions favorable for N₂ fixation in outlet streams.     

Spatial and temporal variation in moss-associated dinitrogen fixation in coniferous- and deciduous-dominated Alaskan boreal forests

Dominant canopy tree species have strong effects on the composition and function of understory species, particularly bryophytes. In boreal forests, bryophytes and their associated microbes are a primary source of ecosystem nitrogen (N) inputs, and an important process regulating ecosystem productivity. We investigated how feather moss-associated N₂-fixation rates and contribution to N budgets vary in time and space among coniferous and broadleaf deciduous forests. We measured N₂-fixation rates using stable isotope (¹⁵N₂) labeling in two moss species (Pleurozium schreberi and Hylocomium splendens) in broadleaf deciduous (Alaska paper birch—Betula neoalaskana) and coniferous (black spruce—Picea mariana) stands near Fairbanks, interior Alaska, from 2013 to 2015. N₂-fixation rates showed substantial inter-annual variation among the 3 years. High N₂-fixation was more strongly associated with high precipitation than air temperature or light availability. Overall, contribution of N₂-fixation to N budgets was greater in spruce than in birch stands. Our results enhance the knowledge of the processes that drive N₂-fixation in boreal forests, which is important for predicting ecosystem consequences of changing forest composition.     

Species richness and nitrogen supply regulate the productivity and respiration of ectomycorrhizal fungi in pure culture

The effects of biodiversity of aboveground organisms have been widely investigated in a range of ecosystems, yet whether similar responses are also seen in belowground microbial communities, such as ectomycorrhizal (EM) fungi, are little understood. We investigated, in vitro, the effects of a gradient of 1–8 species of EM fungi interacting with substratum carbon:nitrogen (C:N) ratio on biomass production and CO₂ efflux. The model experimental systems enabled us to recover and measure biomass of individuals within communities and calculate net selection and complementarity effects. Both biomass and CO₂ efflux increased with species richness particularly under high N concentrations. Moreover, net biodiversity effects were largely positive, driven by both selection and complementarity effects. Our results reveal, in pure culture, the implications of EM species richness on community productivity and C cycling, particularly under high N conditions, and constitute the basis for future experiments under natural conditions.     

Nitrogen fixation by non-legumes in tropical agriculture with special reference to wetland rice

Summary Of the 143 million hectares of cultivated rice land in the world, 75% are planted to wetland rice. Wet or flooded conditions favour biological nitrogen fixation by providing (1) photic-oxic floodwater and surface soil for phototrophic, free-living or symbiotic blue-green algae (BGA), and (2) aphotic-anoxic soil for anaerobic or microaerobic, heterotrophic bacteria. TheAzolla-Anabaena symbiosis can accumulate as much as 200 kg N ha^–1 in biomass. In tropical flooded fields, biomass production from a singleAzolla crop is about 15 t fresh weight ha^–1 or 35 kg N ha^–1. Low tolerance for high temperature, insect damage, phosphorus requirement, and maintenance of inoculum, limit application in the tropics. Basic work on taxonomy, sporulation, and breeding ofAzolla is needed. Although there are many reports of the positive effect of BGA inoculation on rice yield, the mechanisms of yield increase are not known. Efficient ways to increase N₂-fixation by field-grown BGA are not well exploited. Studies on the ecology of floodwater communities are needed to understand the principles of manipulating BGA. Bacteria associated with rice roots and the basal portion of the shoot also fix nitrogen. The system is known as a rhizocoenosis. N₂-fixation in rhizocoenosis in wetland rice is lower than that ofAzolla or BGA. Ways of manipulating this process are not known. Screening rice varieties that greatly stimulate N₂-fixation may be the most efficient way of manipulating the rhizocoenosis. Stimulation of N₂-fixation by bacterial inoculation needs to be quantified.     

CO(2) Exchange and Dinitrogen Fixation of Subterranean Clover in Response to Light Level

Small swards of nodulated subterranean clover plants were grown in pots to a common dry weight under controlled conditions. The rooting medium was a porous calcined clay. All mineral nutrients except nitrogen were supplied daily in solution. Pots then were placed in an assimilation chamber for 3 days for the measurement of net CO₂ exchange at light levels ranging from 0.1 to 2.0 millieinsteins per square meter per second. N₂-fixation (acetylene reduction) of each pot was measured subsequently. H₂-evolution and N₂-fixation were measured for similar treatments in separate experiments using smaller pots.     

Plant functional traits and soil microbial biomass in different vegetation zones on the Loess Plateau

Plant functional traits built the relationships between plant diversity, species composition, and physiology along with the environmental changes, thus influencing soil microbial community. As the sensitivity indicators, soil microbial biomass and plant functional traits responses soil micro-organism and plant characteristics in direct way. Ten plant functional traits of 149 species and soil microbial biomass (carbon, nitrogen, and phosphorus) were analyzed across the different vegetation types (forest, forest-steppe, and steppe) that are divided by environmental gradient (temperature and precipitation), aimed to find the correlations among them. Our results confirmed the greatest values of plant functional traits (except the leaf density and the fine root density) that were distributed in the steppe zone, mainly due to the different mean annual temperature and mean annual precipitation conditions. For different plant growth forms, the plant functional traits were significant differences among the vegetation zones. The advantages of higher rate nutrient cycling, plentiful biomass supplements, and favorite habit conditions lead to the forest-steppe zone with the highest C_mic and N_mic concentrations. The canonical correlation analysis indicated that leaf nitrogen, root nitrogen, and fine root densities were correlated with root exudate and tissue which affected the concentrations of soil organic carbon (SOC) and total nitrogen (N), consequently impacting soil microbial biomass carbon (C_mic) and soil microbial biomass nitrogen (N_mic). Soil is the medium that connects micro-organism and plant root system that influenced leaf nitrogen, root nitrogen, and fine root density of plant functional traits, the concentrations of SOC and total N that plant feedback are consequently influencing C_mic and N_mic.     

Combined effects of CO2 and light on large and small isolates of the unicellular N2-fixing cyanobacterium Crocosphaera watsonii from the western tropical Atlantic Ocean

We examined the combined effects of light and pCO₂ on growth, CO₂-fixation and N₂-fixation rates by strains of the unicellular marine N₂-fixing cyanobacterium Crocosphaera watsonii with small (WH0401) and large (WH0402) cells that were isolated from the western tropical Atlantic Ocean. In low-pCO₂-acclimated cultures (190 ppm) of WH0401, growth, CO₂-fixation and N₂-fixation rates were significantly lower than those in cultures acclimated to higher (present-day ～385 ppm, or future ～750 ppm) pCO₂ treatments. Growth rates were not significantly different, however, in low-pCO₂-acclimated cultures of WH0402 in comparison with higher pCO₂ treatments. Unlike previous reports for C. watsonii (strain WH8501), N₂-fixation rates did not increase further in cultures of WH0401 or WH0402 when acclimated to 750 ppm relative to those maintained at present-day pCO₂. Both light and pCO₂ had a significant negative effect on gross : net N₂-fixation rates in WH0402 and trends were similar in WH0401, implying that retention of fixed N was enhanced under elevated light and pCO₂. These data, along with previously reported results, suggest that C. watsonii may have wide-ranging, strain-specific responses to changing light and pCO₂, emphasizing the need for examining the effects of global change on a range of isolates within this biogeochemically important genus. In general, however, our data suggest that cellular N retention and CO₂-fixation rates of C. watsonii may be positively affected by elevated light and pCO₂ within the next 100 years, potentially increasing trophic transfer efficiency of C and N and thereby facilitating uptake of atmospheric carbon by the marine biota.     

氮素添加和刈割对内蒙古弃耕草地土壤氮矿化的影响

以内蒙古多伦县恢复生态学试验示范研究站弃耕10余年的草地为研究对象,于2006年起分别设置对照、氮素添加、刈割和氮素添加+刈割4种处理,每种处理6次重复,研究弃耕草地氮素添加和刈割对土壤氮矿化的影响,结合土壤理化性质和植被地上生产力的动态变化,分析弃耕草地土壤氮矿化对植被恢复的响应,为当地草地恢复与重建提供理论依据和数据支持。实验结果表明:1氮素添加显著增加了植物地上净初级生产力(ANPP)和土壤无机氮库,与对照相比分别提高115%和196%,同时显著提高了土壤总硝化速率;但是氮素添加对总氨化速率、土壤微生物生物量碳(MBC)、微生物生物量氮(MBN)、微生物生物量碳氮比(MBC/MBN)、微生物呼吸(MR)以及呼吸熵(q CO2)均无显著影响;2总氨化速率和硝化速率对刈割处理的响应均不显著,但是刈割处理显著降低了土壤MR(P0.05);3氮素添加+刈割处理5—7a后,土壤总氨化和硝化速率均无显著变化;但是氮素添加+刈割处理显著增加了ANPP、土壤无机氮库和q CO2,同时显著降低了MBC和MBC/MBN。这说明在弃耕草地适应性管理中,氮素添加可以显著提高草地生产力,但是长期的氮添加对土壤微生物氮的转化是否有利还值得我们进一步研究。