Relationships between soil microbial properties and aboveground stand characteristics of conifer forests in Oregon

Eight forest sites representing a large range of climate, vegetation, and productivity were sampled in a transect across Oregon to study the relationships between aboveground stand characteristics and soil microbial properties. These sites had a range in leaf area index of 0.6 to 16 m² m^–2 and net primary productivity of 0.3 to 14 Mg ha^–1 yr^–1.Measurements of soil and forest floor inorganic N concentrations and in situ net N mineralization, nitrification, denitrification, and soil respiration were made monthly for one year. Microbial biomass C and anaerobic N mineralization, an index of N availability, were also measured. Annual mean concentrations of NH ₄ ⁺ ranged from 37 to 96 mg N kg^–1 in the forest floor and from 1.7 to 10.7 mg N kg^–1 in the mineral soil. Concentrations of NO ₃ ^– were low ( < 1 mg N kg^–1) at all sites. Net N mineralization and nitrification, as measured by the buried bag technique, were low on most sites and denitrification was not detected at any site. Available N varied from 17 to 101 mg N kg^–1, microbial biomass C ranged from 190 to 1230 mg Ckg^–1, and soil respiration rates varied from 1.3 to 49 mg C kg^–1 day^–1 across these sites. Seasonal peaks in NH ₄ ⁺ concentrations and soil respiration rates were usually observed in the spring and fall.The soils data were positively correlated with several aboveground variables, including leaf area index and net primary productivity, and the near infrared-to-red reflectance ratio obtained from the airborne simulator of the Thematic Mapper satellite. The data suggest that close relationships between aboveground productivity and soil microbial processes exist in forests approaching semi-equilibrium conditions.Abbreviations IR infrared - LAI leaf area index - k _c proportion of microbial biomass C mineralized to CO₂ - NPP net primary productivity - TM Thematic Mapper     

Effects of a clearcut on the net rates of nitrification and N mineralization in a northern hardwood forest,Catskill Mountains,New York,USA

The Catskill Mountains of southeastern New York receive among the highest rates of atmospheric nitrogen (N) deposition in eastern North America, and ecosystems in the region may be sensitive to human disturbances that affect the N cycle. We studied the effects of a clearcut in a northern hardwood forest within a 24-ha Catskill watershed on the net rates of N mineralization and nitrification in soil plots during 6 years (1994–1999) that encompassed 3-year pre- and post-harvesting periods. Despite stream NO₃^– concentrations that increased by more than 1400 mol l^–1 within 5 months after the clearcut, and three measures of NO₃^– availability in soil that increased 6- to 8-fold during the 1^st year after harvest, the net rates of N mineralization and nitrification as measured by in situ incubation in the soil remained unchanged. The net N-mineralization rate in O-horizon soil was 1– 2 mg N kg^–1 day^–1 and the net nitrification rate was about 1 mg N kg^–1 day^–1, and rates in B-horizon soil were only one-fifth to one-tenth those of the O-horizon. These rates were obtained in single 625 m² plots in the clearcut watershed and reference area, and were confirmed by rate measurements at 6 plots in 1999 that showed little difference in N-mineralization and nitrification rates between the treatment and reference areas. Soil temperature increased 1 ± 0.8 °C in a clearcut study plot relative to a reference plot during the post-harvest period, and soil moisture in the clearcut plot was indistinguishable from that in the reference plot. These results are contrary to the initial hypothesis that the clearcut would cause net rates of these N-cycling processes to increase sharply. The in situ incubation method used in this study isolated the samples from ambient roots and thereby prevented plant N uptake; therefore, the increases in stream NO₃^– concentrations and export following harvest largely reflect diminished uptake. Changes in temperature and moisture after the clearcut were insufficient to measurably affect the net rates of N mineralization and nitrification in the absence of plant uptake. Soil acidification resulting from the harvest may have acted in part to inhibit the rates of these processes. The US Governments right to retain a non-exclusive, royalty-free license in and to any copyright is acknowledged.     

Net nitrogen mineralization in soils under six indigenous tree species,an abandoned pasture and a secondary forest in the atlantic lowlands of costa rica

Nitrogen mineralization, nitrification potentials, pH, total N, C, extractable P and cations were measured in soils under 4-year-old, mono-specific stands of six fast-growing, native tree species, an abandoned pasture, and a 20-year-old secondary forest, as part of a study on the use of indigenous tree species for rehabilitation of soil fertility on degraded pastures at the La Selva Biological Station in the Atlantic humid lowlands of Costa Rica. Soil net nitrification potential rates were higher under two N-fixing, leguminous species,Stryphnodendron microstachyum Poepp. et Endl. (1.1–1.9 mg kg^–1 day^–1) andDalbergia tucurensis Donn. Smith (0.7–1.5 mg kg^–1 day^–1), than under the non-N-fixing trees in the plantation,Vochysia guatemalesis Don. Sm.,Vochysia ferruginea Mart,Dipteryx panamensis (Pittier) Record and Mell andHyeronima alchorneoides Fr. Allemao (0.2–0.8 mg kg^–1 day^–1). Values under the N-fixing trees were comparable to those found in secondary forest. There were no statistically significant differences in soil total N or in other nurtients between the species. Results of pH measurements done before and after incubation did not show any clear evidence of a pH drop attributable to nitrification.     

Effects of understory removal, N fertilization, and litter layer removal on soil N cycling in a 13-year-old white spruce plantation infested with Canada bluejoint grass

Canada bluejoint grass [Calamagrostis canadensis (Michx.) Beauv., referred to as bluejoint below] is a competitive understory species widely distributed in the boreal region in North America and builds up a thick litter layer that alters the soil surface microclimate in heavily infested sites. This study examined the effects of understory removal, N fertilization, and litter layer removal on litter decomposition, soil microbial biomass N (MBN), and net N mineralization and nitrification rates in LFH (the sum of organic horizons of litter, partially decomposed litter and humus on the soil surface) and mineral soil (0–10 cm) in a 13-year-old white spruce [Picea glauca (Moench.) Voss] plantation infested with bluejoint in Alberta, Canada. Removal of the understory vegetation and the litter layer together significantly increased soil temperature at 10 cm below the mineral soil surface by 1.7 and 1.3°C in summer 2003 and 2004, respectively, resulting in increased net N mineralization (by 1.09 and 0.14 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004) and net nitrification rates (by 0.10 and 0.20 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004). When the understory vegetation was intact, nitrification might have been limited by NH₄ ⁺ availability due to competition for N from bluejoint and other understory species. Litter layer removal increased litter decomposition rate (percentage mass loss per month) from 2.6 to 3.0% after 15 months of incubation. Nitrogen fertilization did not show consistent effects on soil MBN, but increased net N mineralization and nitrification rates as well as available N concentrations in the soil. Clearly, understory removal combined with N fertilization was most effective in increasing rates of litter decomposition, net N mineralization and nitrification, and soil N availability. The management of understory vegetation dominated by bluejoint in the boreal region should consider the strong effects of understory competition and the accumulated litter layer on soil N cycling and the implications for forest management.     

Quantitative traits associated with adaptation of three barley (Hordeum vulgare L) cultivars to suboptimal iron supply

A laboratory method was developed that allows determination of in situ net nitrification with high sensitivity and at high temporal resolution. Nitrate in soils is quantitatively converted into nitrous oxide under strictly anaerobic conditions in the presence of 10 kPa acetylene by the soil endogenous denitrifier population, with the N₂O detected by a gas chromatograph equipped with a ⁶³Ni electron capture detector. Thus, even low net nitrification rates, i.e. small net increases in soil nitrate concentrations can easily be detected. Comparison of results using this method with results obtained using the classical in situ incubation method (buried bag soil incubation) revealed excellent agreement. Application of the new method allowed both determination of the seasonal pattern of net nitrification as well as correlation analysis between in situ NO and N₂O flux rates and in situ net nitrification rates of the forest soils studied. Regardless of the forest site studied (spruce, spruce limed, beech), and during each year of a 3 years period (1995–1997), net nitrification varied strongly with season and was least during winter and greatest during summer. The long-term annual, mean rate of net nitrification for the untreated spruce site, the limed spruce site and the beech site were 1.54 ± 0.27 mg N kg^–1 sdw d^–1, 1.92 ± 0.23 mg N kg^–1 sdw d^–1 and 1.31 ± 0.23 mg N kg^–1 sdw d^–1, respectively. In situ rates of nitrification and NO and N₂O emission were strongly correlated for all sites suggesting that nitrification was the dominate source of NO as well as N₂O.     

晋西北不同年限小叶锦鸡儿灌丛土壤氮矿化和硝化作用

利用PVC管顶盖埋管法研究了晋西北黄土高原区小叶锦鸡儿人工灌丛不同定植年限(5,10,20,30,40a)土壤氮矿化与硝化速率的动态和净矿化与硝化总量。结果表明,⑴小叶锦鸡儿灌丛土壤无机氮主要以NO_-~3-N形式存在,不同生长年限相同月份的土壤硝态氮(NO-3-N)含量分别是铵态氮(NH+4-N)含量的1.5—15.4倍;⑵土壤氮素硝化速率和矿化速率随生长年限延长而加快,30年生时达到高峰,数值达40.2,44.1 mg m~(-2)d~(-1)。从季节性变化看,7—8月份是硝化速率和矿化速率快速增长期,30年生小叶锦鸡儿灌丛土壤硝化速率和矿化速率分别达到86.9,93.1 mg m~(-2)d~(-1),显著高于其它生长年限(P0.05);(3)土壤氮素硝化与矿化总量同样随小叶锦鸡儿生长年限延长而增加,30年生时达到最高,与5年生相比,分别增加了3.7和3.1倍。(4)5—10月份小叶锦鸡儿生长期内,各年限土壤全氮量的2.3%被矿化成无机氮,其中87%最终被转化成NO-3-N形式存在于土体中。     

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.     

大兴安岭北部天然针叶林土壤氮矿化特征

采用顶盖埋管法对大兴安岭地区天然针叶林(樟子松林、樟子松-兴安落叶松混交林和兴安落叶松林)土壤铵态氮(NH~+_4-N)、硝态氮(NO~-_3-N)、净氮矿化速率进行研究,并探索土壤理化性质与氮矿化之间的相关性,为大兴安岭地区森林生态系统土壤养分管理及森林经营提供帮助。结果表明:观测期内(5—10月)3种林型土壤无机氮变化范围为31.51—70.42 mg/kg,以NH~+_4-N形式存在为主,占比达90%以上,且与纯林相比混交林土壤无机氮含量较高。3种林型土壤净氮矿化、净氨化、净硝化速率月变化趋势呈V型,7、8月表现为负值,其他月份为正值。净氮矿化速率变化范围樟子松林为-0.54—1.28 mg kg~(-1) d~(-1)、樟子松-兴安落叶松混交林为-0.13—0.55 mg kg~(-1) d~(-1)、兴安落叶松林为-0.80—1.05 mg kg~(-1) d~(-1)。土壤净氨化过程在土壤氮矿化中占主要地位,占比达60%以上。3种林型土壤净氮矿化、净氨化及净硝化速率垂直差异显著,0—10 cm土层矿化作用明显高于10—20 cm土层(P0.05)。土壤氮矿化速率与土壤含水量、土壤有机碳含量、土壤C/N、枯落物全氮含量和枯落物C/N均存在显著相关性。不同类型的森林土壤及枯落物的质量也存在差异,进而影响土壤氮矿化特征。     

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.     

Net mineralization and nitrification rates in a clay soil measured and predicted in permanent grassland from soil temperature and moisture content

Summary Net mineralization of N and net nitrification in field-moist clay soils (Evesham-Kingston series) from arable and grassland sites were measured in laboratory incubation experiments at 4, 10 and 20°C. Three depth fractions to 30 cm were used. Nitrate accumulated at all temperatures except when the soil was very dry (=0.13 cm³ cm^–3). Exchangeable NH₄-ions declined during the first 24 h and thereafter remained low. Net mineralization and net nitrification approximated to zero-order reactions after 24 h, with Q₁₀ values generally <1.6. The effect of temperature on both processes was linear although some results conformed to an Arrhenius-type relationship. The dependence of net mineralization and net nitrification in the field soil on soil temperature (10 cm depth) and moisture (0–15, 15–25, 25–35 cm depths) was modelled using the laboratory incubation data. An annual net mineralization of 350 kg N ha^–1 and net nitrification of 346 kg N ha^–1 were predicted between September 1980 and August 1981. The model probably overstressed the effect of soil moisture relative to soil temperature.     

Mineral nitrogen and microbial dynamics in the forest floor of clearcut or partially harvested successional boreal forest stands

The effects of clearcut and partial harvesting of early-seral trembling aspen plots were compared to conventional clearcut harvesting in mid-seral mixedwood and late-seral conifer plots. Twice a year, for three consecutive years, we assessed mineral N and microbial dynamics in the forest floor of these plots to test three hypotheses related to the higher litter quality of aspen leaves and to the sustained inputs of available C on partially harvested plots: (1) the post-clearcutting mineral N flush and the net [(NO₃^–): (NO₃^– + NH₄⁺)] production ratio (RNI) are higher in aspen plots than in black spruce plots, with intermediate values occurring in mixedwood plots; (2) net N mineralization rates in aspen plots are higher in spring than in autumn; and (3) compared to clearcutting, partial harvesting reduces potential ammonification and nitrification rates. Initial NH₄⁺ and NO₃^– concentrations respectively ranged between 1.7–4.4 and 0.2–1.5 g N kg^–1 N_total, net ammonification and nitrification rates (30 d incubations) respectively ranged between 5.3–17.8 and 0.1–27.6 g N kg^–1 N_total, basal respiration ranged between 20.9–38.9 mg CO₂-C kg^–1 h^–1, and microbial biomass ranged between 6.1–8.7 g C_mic kg^–1. Although clearcutting increased NO₃^– concentrations in aspen plots, the balance of our results did not support our first hypothesis, because NH₄⁺ concentrations increased in conifer plots only, potential ammonification was unaffected by clearcutting, potential nitrification increased in mixedwood plots only, and RNI increased in all plots. In each seral stage, basal respiration, microbial biomass, and metabolic quotient either increased or were unaffected by clearcutting, suggesting that increases in RNI after disturbance were not related to lower microbial immobilisation of NO₃^– due to lower available C. Forest floors in mid-seral mixedwood plots exhibited a distinct combination of mineral N and microbial properties, suggesting that the functional richness of the forest is enhanced not only by the number of species, but also by the diversity of assemblages that are present. Results supported our second hypothesis and showed, furthermore, that net N mineralization in conifer stands is greater in autumn than in spring. Partial harvesting in aspen stands resulted in lower potential mineralization of N and lower RNI, compared to clearcutting. Further lysimetry studies are needed to confirm whether partial harvesting mitigates NO₃^– leaching following disturbance.     

Net nitrogen mineralization and net nitrification along a tropical forest-to-pasture chronosequence

Soil inorganic nitrogen pools, net mineralization and net nitrification rates were compared during the dry season along a chronosequence of upland (terra firme) forest, 3-, 9- and 20-year-old pastures in the western Brazilian Amazon Basin state of Rondônia to investigate the influence of forest conversion to pasture on soil nitrogen cycles. Surface soil (0 to 10 cm) from forest had larger extractable inorganic nitrogen pools than pasture soils. In the forest, NO ₃ ^– pools equaled or exceeded NH ₄ ⁺ pools, while pasture inorganic N pools consisted almost exclusively of NH ₄ ⁺ . Rates of net N mineralization and net nitrification in seven -day laboratory incubations were higher in the seven - day forest than in the pastures. Net N mineralization rates did not differ significantly among different-aged pastures, but net nitrification rates were significantly lower in the 20-year-old pasture. Higher net N mineralization and net nitrification rates were measured in laboratory and in situ incubations of sieved soil, compared with in situ incubations of intact soil cores. Rates calculated in seven-day incubations were higher than determined by longer incubations. Sieving may increase N mineralization and/or decrease N immobilization compared with intact cores. We concluded that 7-day laboratory incubation of sieved soil was the most useful index for comparing N availability across the chronosequence of forest and pasture sites. High net nitrification rates in forest soils suggest a potential for NO ₃ ^– losses either through leaching or gaseous emissions.     

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 (r ² = 0.37, P 0.01). This pattern in the field corresponded to the pattern shown in the laboratory assays with increasing monoterpene additions.     

Factors affecting nitrogen dynamics in a semiarid woodland (Dry Chaco,Argentina)

In an effort to elucidate the factors affecting soil N dynamics in the Dry Chaco ecosystem, soil respiration and microbial biomass N were measured for one year underneath 5 vegetation types: a leguminous tree (Prosopis flexuosa DC), a non-leguminous tree (Aspidosperma quebracho-blanco Schlecht.), a non leguminous shrub (Larrea spp.), the open interspaces, and a pure grassland. Ammonifier and nitrifier densities and N content in litter were also measured in some cases. Results were compared with previously reported N mineralization rates and soil fertility.During the dry season microbial biomass N and net N mineralization were low, while accretion of easily mineralizable C occurred (estimated through soil respiration rates in lab under controlled temperature and moisture). With the onset of rain, microbial biomass N and N mineralization increased markedly, resulting in a decrease in easily mineralizable C. Throughout the wet season N mineralization varied with soil moisture while microbial biomass N remained consistently high. Mean values of immobilized N in this ecosystem were high (20–140 mg kg^–1), of about the same order of magnitude as accumulated net N mineralization (50–150 mg kg^–1 yr^–1). Microbial decay in the dry season, considered as a source of easily mineralizable N, accounted for only 40% of gross N mineralization increase at the beginning of the wet season. Ammonifier densities correlated significantly with soil moisture and N mineralization, but nitrifiers did not.The highest values of total N, N mineralization, inorganic N, microbial biomass N, nitrifier densities, N content in litter, total organic C and easily mineralizable C were found under Prosopis and the lowest values under shrubs and the interspaces. The main differences between tree species were in N mineralization at the beginning of the wet season, in total and inorganic N pools, and in nitrifier densities; all of which were significantly lower under Aspidosperma than under Prosopis.N mineralization in the pure grassland was very low despite high values of total N and C sources. Although N immobilized in microbial biomass was similarly high under Aspidosperma, Prosopis and the pure grassland, net N mineralization rates were quite different.     

植被类型与坡位对喀斯特土壤氮转化速率的影响

土壤氮素转化对于植物氮素营养具有重要作用,尤其是对于受氮素限制的喀斯特退化生态系统。选取植被恢复过程中4种典型喀斯特植被类型(草丛、灌丛、次生林、原生林)和3个坡位(上、中、下坡位)表层土壤(0—15cm)为对象,利用室内培养的方法,研究不同植被类型和坡位下土壤氮素养分与氮转化速率(氮净矿化率、净硝化率和净氨化率)的特征及其影响因素。结果表明,植被类型对土壤硝态氮含量、无机氮含量、氮净矿化率、净硝化率和净氨化率均有显著影响(P0.01),即随着植被的正向演替(草丛—灌丛—次生林—原生林),土壤硝态氮含量、无机氮含量、土壤氮净矿化速率和净硝化速率整体上呈增加趋势,而坡位以及坡位与植被类型的交互作用对上述土壤氮素指标无显著影响(P0.05)。冗余分析结果表明凋落物氮含量、凋落物C∶N比和硝态氮含量对土壤氮转化速率有显著影响,其中凋落物氮含量是影响土壤氮转化速率的主要因子(F=35.634,P=0.002)。可见,尽管坡位影响喀斯特水土再分配过程,但植被类型决定的凋落物质量(如凋落物氮含量等)对喀斯特土壤氮素转化速率的作用更为重要。因此,在喀斯特退化生态系统植被恢复初期,应注重植被群落的优化配置(如引入豆科植物)和土壤质量的改善(如降低土壤C∶N),促进土壤氮素转化及氮素的有效供给。     

Land-use change effects on soil C and N transformations in soils of high N status: comparisons under indigenous forest, pasture and pine plantation

Globally, land-use change is occurring rapidly, and impacts on biogeochemical cycling may be influenced by previous land uses. We examined differences in soil C and N cycling during long-term laboratory incubations for the following land-use sequence: indigenous forest (soil age = 1800 yr); 70-year-old pasture planted after forest clearance; 22-year-old pine (Pinus radiata) planted into pasture. No N fertilizer had been applied but the pasture contained N-fixing legumes. The sites were adjacent and received 3–6 kg ha^–1 yr^–1volcanic N in rain; NO₃ ^–-N leaching losses to streamwater were 5–21 kg ha^–1 yr^–1, and followed the order forest < pasture = pine. Soil C concentration in 0–10 cm mineral soil followed the order: pasture > pine = forest, and total N: pasture > pine > forest. Nitrogen mineralization followed the order: pasture > pine > forest for mineral soil, and was weakly related to C mineralization. Based on radiocarbon data, the indigenous forest 0–10 cm soil contained more pre-bomb C than the other soils, partly as a result of microbial processing of recent C in the surface litter layer. Heterotrophic activity appeared to be somewhat N limited in the indigenous forest soil, and gross nitrification was delayed. In contrast, the pasture soil was rich in labile N arising from N fixation by clover, and net nitrification occurred readily. Gross N cycling rates in the pine mineral soil (per unit N) were similar to those under pasture, reflecting the legacy of N inputs by the previous pasture. Change in land use from indigenous forest to pasture and pine resulted in increased gross nitrification, net nitrification and thence leaching of NO₃ ^–-N.     

Ungulate stimulation of nitrogen cycling and retention in Yellowstone Park grasslands

We studied how ungulates and a large variation in site conditions influenced grassland nitrogen (N) dynamics in Yellowstone National Park. In contrast to most grassland N studies that have examined one or two soil N processes, we investigated four rates, net N mineralization, nitrification, denitrification, and inorganic N leaching, at seven paired sites inside and outside long-term (33+ year) exclosures. Our focus was how N fluxes were related to one another among highly variable grasslands and how grazers influenced those relationships. In addition, we examined variation in soil δ¹⁵N among grasslands and the relationships between soil ¹⁵N abundance and N processes. Previously, ungulates were reported to facilitate net N mineralization across variable Yellowstone grasslands and denitrification at mesic sites. In this study, we found that herbivores also promoted nitrification among diverse grasslands. Furthermore, net N mineralization, nitrification, and denitrification (kg N ha^–1 year^–1, each variable) were postively and linearly related to one another among all grasslands (grazed and fenced), and grazers reduced the nitrification/net N mineralization and denitrification/net N mineralization ratios, indicating that ungulates inhibited the proportion of available NH₄ ⁺ that was nitrified and denitrified. There was no relationship between net N mineralization or nitrification with leaching (indexed by inorganic N adsorbed to resin buried at the bottom of rooting zones) and leaching was unaffected by grazers. Soil δ¹⁵N was positively and linearly related to in situ net N mineralization and nitrification in ungrazed grasslands; however, there was no relationship between isotopic composition of N and those rates among grazed grasslands. The results suggested that grazers simultaneously increased N availability (stimulated net N mineralization and nitrification per unit area) and N conservation (reduced N loss from the soil per unit net N mineralization) in Yellowstone grasslands. Grazers promoted N retention by stimulating microbial productivity, probably caused by herbivores promoting labile soil C. Process-level evidence for N retention by grazers was supported by soil δ¹⁵N data. Grazed grassland with high rates of N cycling had substantially lower soil δ¹⁵N relative to values expected for ungrazed grassland with comparable net N mineralization and nitrification rates. These soil ¹⁵N results suggest that ungulates inhibited N loss at those sites. Such documented evidence for consumer control of N availability to plants, microbial productivity, and N retention in Yellowstone Park is further testimony for the widespread regulation of grassland processes by large herbivores. Received: 5 May 1999 / Accepted: 1 November 1999     

Soil-mixing effects on inorganic nitrogen production and consumption in forest and shrubland soils

Soils that are physically disturbed are often reported to show net nitrification and NO₃⁻ loss. To investigate the response of soil N cycling rates to soil mixing, we assayed gross rates of mineralization, nitrification, NH₄⁺ consumption, and NO₃⁻ consumption in a suite of soils from eleven woody plant communities in Oregon, New Mexico, and Utah. Results suggest that the common response of net NO₃⁻ flux from disturbed soils is not a straightforward response of increased gross nitrification, but instead may be due to the balance of several factors. While mineralization and NH₄⁺ assimilation were higher in mixed than intact cores, NO₃⁻ consumption declined. Mean net nitrification was 0.12 mg N kg⁻¹ d⁻¹ in disturbed cores, which was significantly higher than in intact cores (−0.19 mg N kg⁻¹ d⁻¹). However, higher net nitrification rates in disturbed soils were due to the suppression of NO₃⁻ consumption, rather than an increase in nitrification. Our results suggest that at least in the short term, disturbance may significantly increase NO₃⁻ flux at the ecosystem level, and that N cycling rates measured in core studies employing mixed soils may not be representative of rates in undisturbed soils.     

Nitrogen mineralization in heathland ecosystems dominated by different plant species

Net N mineralization rates were measured in heathlands still dominated by ericaceous dwarf shrubs (Calluna vulgaris or Erica tetralix) and in heathlands that have become dominated by grasses (Molinia caerulea or Deschampsia flexuosa). Net N mineralization was measuredin situ by sequential soil incubations during the year. In the wet area (gravimetric soil moisture content 74–130%), the net N mineralization rates were 4.4 g N m^–2 yr^–1 in the Erica soil and 7.8 g N m^–2 yr^–1 in the Molinia soil. The net nitrification rate was negligibly slow in either soil. In the dry area (gravimetric soil moisture content 7–38%), net N mineralization rates were 6.2 g N M-2 yr^–1 in the Calluna soil, 10.9 g N m^–2 yr^–1 in the Molinia soil and 12.6 g N m^–2 yr^–1 in the Deschampsia soil. The Calluna soil was consistently drier throughout the year, which may partly explain its slower mineralization rate. Net nitrification was 0.3 g N m^–2 yr^–1 in the Calluna soil, 3.6 g N m^–2 yr^–1 in the Molinia soil and 5.4 g N m^–2 yr^–1 in the Deschampsia soil. The net nitrification rate increased proportionally with the net N mineralization rate suggesting ammonium availability may control nitrification rates in these soils. In the dry area, the faster net N mineralization rates in sites dominated by grasses than in the site dominated by Calluna may be explained by the greater amounts of organic N in the soil of sites dominated by grasses. In both areas, however, the net amount of N mineralized per gram total soil N was greater in sites dominated by Molinia or Deschampsia than in sites dominated by Calluna or Erica. This suggests that in heathlands invaded by grasses the quality of the soil organic matter may be increased resulting in more rapid rates of soil N cycling.     

Soil nitrogen mineralization in plantations ofJuglans nigra interplanted with actinorhizalElaeagnus umbellata orAlnus glutinosa

Nitrogen mineralization rates were estimated in 19-year-old interplantings of black walnut (Juglans nigra L.) with dinitrogen fixing autumn-olive (Elaeagnus umbellata Thunb.) or black alder (Alnus glutinosa L. Gaertn.) and in pure walnut plantings at two locations in Illinois USA. N mineralization rates were measured repeatedly over a one year period usingin situ incubations of soil cores in oxygen-permeable polyethylene bags at 0–10 and 10–20 cm soil depths, and also by burying mixed-bed ion-exchange resin in soil. Mineralization rates were highest in summer and in plots containing actinorhizal Elaeagnus and Alnus in contrast with pure walnut plots. Elaeagnus plots at one location yielded 236 kg of mineral N ha^–1 yr^–1 in the upper 20 cm of soil, a value higher than previously reported for temperate decidous forest soils in North America. The highest mean plot values for N mineralization in soil at a location were 185 kg ha^–1 yr^–1 for Alnus interplantings and 90 kg ha^–1 yr^–1 for pure walnut plots. Plots which had high N mineralization rates also had the largest walnut trees. Despite low pH (4.1 and 6.5) and low extractable P concentrations (1.4 and 0.7 mg kg^–1 dry mass) at the two locations, nitrification occurred in all plots throughout the growing season. NO ₃ ^– –N was the major form of mineralized N in soil in the actinorhizal interplantings, with NH ₄ ⁺ –N being the major form of mineral N in control plots. Walnut size was highly correlated with soil nitrogen mineralization, particularly soil NO ₃ ^– –N production in a plot.