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     

Microbial Nitrogen and Sulfur Cycles at the Gypsum Dunes of White Sands National Monument,New Mexico

The White Sands National Monument from New Mexico (U.S.A) contains one of the largest known gypsum dune fields with unique, rapidly migrating, arid, evaporitic habitats. Deposits from dune sides and interdune areas were collected in order to determine the characteristics of microbial habitat and communities through mineral assemblages, microbial pigments along with investigations of nitrogen and sulfur cycles. The most abundant pigments, scytonemin and carotenoids, were common UV protective pigments. Predominance of nitrite and nitrate over ammonium nitrogen (2.16: 1) implies that nitrification processes might be important in this ecosystem. Ammonium oxidizers from groups of β-, γ-proteobacteria and archaea were detected in all deposits, thereby indicating microbial involvement in nitrification. Additionally, denitrifying organisms with nirS and nirK genes were also present in most of the analyzed samples. The presence of trace carbonate mineral phases in association with biofilm implies possible microbial sulfate reduction. Microbes with metabolic abilities for sulfur cycling (i.e., dissimilatory sulfite reducers, purple sulfur bacteria, green sulfur and non-sulfur bacteria, and organisms with the APS enzyme) were identified in all samples. These particular organisms have the ability to reduce sulfate and to re-oxidize reduced sulfur compounds back to sulfate.     

Nitrogen cycling in dense plantings of hybrid poplar and black alder

Summary Nitrogen cycling was studied during the third growing season in pure and mixed plantings (33×33 cm spacing) of hybrid poplar and black alder in southeastern Canada. After 3 years, hybrid poplar growth and N content of living tissues in a plot and of individual hybrid poplar plants increased with the proportion of black alder in a planting. No differences were detected among N contents of individual alder plants regardless of plot treatment. Black alder allocated a larger portion of its N to roots than hybrid poplar. Symbiotic nitrogen fixation was estimated to account for 80% of the nitrogen in aboveground alder tissues in the pure treatment using natural¹⁵N dilution. N return in leaf litter was estimated to be 70kg ha^–1 in the pure alder treatment and decreased to a minimum of 20 kg ha^–1 in the pure hybrid poplar plots. No difference was detected among treatments for throughfall N content. Nitrogen concentration in roots and leaf litterfall of black alder was higher than hybrid poplar. Significant soil N accretion occurred in mixed plantings containing two alders to one poplar and pure black alder plantings. Nitrogen availability (NO₃–N) increased with the amount of black alder in a plot. Results suggest that the early increase in nitrogen accumulation of hybrid poplar in mixed treatments can be attributed to an increase of total soil N availability resulting from the input of large amounts of N from easily mineralizable alder tissue.     

Plant and soil nitrogen dynamics in mediterranean grasslands: a comparison of annual and perennial grasses

Summary The predominance of annual species in the rangelands of southwestern Spain is not due only to climatic factors but is also strongly influenced by grazing management. Manipulating the grazing system in an experimental plot gave a vegetation structure with patches of annual grasses (mainly Vulpia ssp. and Bromus hordeaceus) and patches of perennial grasses (mainly Phalaris aquatica). This vegetation change allowed us to test the hypothesis that life-cycle differences between annual and perennial grasses affect soil nitrogen availability and plant uptake. Nitrogen availability, measured by in situ incubation, and nitrogen uptake were measured through the growing period (October to June). Amounts of in situ mineralized nitrogen over the whole growth phase were more important for soils supporting perennials (37 ppm) than for soils supporting annuals (27 ppm). The difference between the mineral nitrogen produced in situ and the mineral nitrogen accumulated during the same time in the soil allowed an estimation of the maximum mineral nitrogen quantity which can be taken up by the vegetation during each incubation period. The quantities accumulated over the year were 47 and 38 ppm (or 103 and 83 kg/ha) for soils supporting perennials and annuals respectively. For the same period, amounts of nitrogen immobilized in biomass production were 90 and 70 kg/ha for perennials and annuals respectively. During the autumn, a large proportion of mineral nitrogen was leached from soils supporting annual plants which had only just commenced germination. By contrast, the ability to use mineral nitrogen as soon as autumn rains occurred gave a competitive advantage to the perennial species, but only if they were protected from grazing during this period. The higher mineralization and use of this nitrogen reserve by perennials indicates that they made more efficient use of nitrogen resources than annuals, and validate the initial hypothesis.     

Vegetation,biomass and productivity of seral grasslands of Cherrapunji in north-east India

This study deals with the composition of vegetation, biomass and productivity of four different grassland types at Cherrapunji, India. Varied levels of degradation of the climax mexed evergreen forest are represented by these grasslands. Where the soil profile is deep, the grasslands are maintained through frequent fire. At each site, belowground production is more than double aboveground production. Annual turnover rates for the belowground biomass in these grasslands are more dynamical than their temperature counterparts. These grasslands are maintained under high stress conditions of thin soil cover, highly leached nutrient-deficient soils, frequent fires, and low soil moisture retention in spite of a high annual rainfall of 10372 mm.     

Nitrogen budget under rotational bush fallow agriculture (jhum) at higher elevations of Meghalaya in north-eastern India

Summary The nitrogen budge of rotational bush fallow agriculture (jhum) was investigated at higher elevations of Meghalaya in north-eastern India under 15, 10 and 5 year fallow cycles (the intervening fallow period between one or two croppings on the same site). Nitrogen depletion was affected by initial stocks in the soil and vegetation compartment at the time of slash and burn as well as the rate at which this was lost during the subsequent land use. While nitrogen losses due to the burn was more severe under longer cycles compared to the 5 year cycle the losses through sediment and water was more under a 15 year cycle compared to 10 and 5 year cycles. Transfer of nitrogen from soil to the weed biomass increased with shortening of the fallow cycle. The positive role of weeds in conservation of nitrogen in their biomass and subsequent release through organic manure into the agriculture system has been highlighed. Under a short fallow cycle of 5 years, considered on a time scale of 15 years, the soil nitrogen was depleted to a very low level compared to a 15 year cycle, suggesting that a 5 year cycle as prevalent today is not viable from the point of view of nitrogen economy.     

Nitrogen cycling in tropical and temperate savannas

Savannas are the most common vegetation type in the tropics and subtropics, ranging in physiognomy from grasslands with scattered woody plants to woodlands with heterogeneous grass cover. Productivity and organic matter turnover in savannas are controlled by interactions between water and nutrient availability, and this basic environmental structure is modified by fire frequency and land management practices. We compared temperate and tropical savannas in order to understand the strength of nitrogen (N) limitation of productivity. American tropical and temperate savannas are N limited systems, and the N cycle differs according to the woody plant density, fire frequency, land use change, N deposition and N fixation. Grazing and conversion to pasture have been the predominant land-use changes in most savannas. In the Cerrado and the Llanos tropical savannas, intensified use of fire for pasture management is leading to decreased woody plant density. Oppositely, in the Chaco and North American temperate savannas, fire suppression and grazing are leading to increases in woody density. In addition, the higher soil P availability in the Gran Chaco and the higher N deposition in North American savannas may be contributing to increases of N cycling and net productivity rates. Some aspects of the N budget for savannas of the American continent are still unclear and require further analysis to determine rates of N fixation, and to understand how spatial and temporal soil heterogeneity control N fluxes through soil solution and into streams.     

Nutrient cycling and nitrogen mineralization in eucalypt forests of south-eastern Australia

Summary Nutrient pools in litter and soil and the major nutrient transfers and additions in rainfall, throughfall and litterfall were measured in eight mature, undisturbed eucalypt forests covering a range of species, climate, productivity and soil type. Litterfall is the major pathway for the return of N, P, Ca and usually Mg, to the soil. The forests covered almost the range of litterfall reported for eucalypt forests and, in conjunction with published data, litterfall was strongly related to climatic variables. Extractable P in the soil and P concentrations in litter and litterfall were significantly higher in two sub-alpine forests (Eucalyptus pauciflora andE. delegatensis) than in all other forests. In general, nutrient turnover, particularly N turnover, was related to the rate of organic matter turnover. Rates of organic matter turnover in these forests and in other studies of eucalypts were correlated with climatic conditions using the simple climatic scalar developed by Vitousek. Nitrogen turnover, especially that proportion cycling via leaf litterfall is primarily a function of organic matter turnover, but litter quality appears also to have an influence.     

Grazing-induced changes in plant composition affect litter quality and nutrient cycling in flooding Pampa grasslands

Changes in plant community composition induced by vertebrate grazers have been found to either accelerate or slow C and nutrient cycling in soil. This variation may reflect the differential effects of grazing-promoted (G+) plant species on overall litter quality and decomposition processes. Further, site conditions associated with prior grazing history are expected to influence litter decay and nutrient turnover. We studied how grazing-induced changes in plant life forms and species identity modified the quality of litter inputs to soil, decomposition rate and nutrient release in a flooding Pampa grassland, Argentina. Litter from G+ forbs and grasses (two species each) and grazing-reduced (G−) grasses (two species) was incubated in long-term grazed and ungrazed sites. G+ species, overall, showed higher rates of decomposition and N and P release from litter. However, this pattern was primarily driven by the low-growing, high litter-quality forbs included among G+ species. Forbs decomposed and released nutrients faster than either G+ or G− grasses. While no consistent differences between G+ and G− grasses were observed, patterns of grass litter decay and nutrient release corresponded with interspecific differences in phenology and photosynthetic pathway. Litter decomposition, N release and soil N availability were higher in the grazed site, irrespective of species litter type. Our results contradict the notion that grazing, by reducing more palatable species and promoting less palatable ones, should decrease nutrient cycling from litter. Plant tissue quality and palatability may not unequivocally link patterns of grazing resistance and litter decomposability within a community, especially where grazing causes major shifts in life form composition. Thus, plant functional groups defined by species’ “responses” to grazing may only partially overlap with functional groups based on species “effects” on C and nutrient cycling.     

Nitrogen deposition,distribution and cycling in a subalpine spruce-fir forest in the Adirondacks,New York,USA

Nitrogen inputs, fluxes, internal generation and consumption, and outputs were monitored in a subalpine spruce-fir forest at approximately 1000-m elevation on Whiteface Mountain in the Adirondacks of New York, USA. Nitrogen in precipitation, cloudwater and dry deposition was collected on an event basis and quantified as an input. Throughfall, stemflow, litterfall and soil water were measured to determine fluxes within the forest. Nitrogen mineralization in the forest floor was estimated to determine internal sources of available N. Lower mineral horizon soil water was used to estimate output from the ecosystem. Vegetation and soil N pools were determined.During four years of continuous monitoring, an average of 16 kg N ha^–1 yr^–1 was delivered to the forest canopy as precipitation, cloudwater and dry deposition from the atmosphere. Approximately 30% of the input was retained by the canopy. Canopy retention is likely the result of both foliar uptake and immobilization by bark, foliage and microorganisms. Approximately 40 kg of N was made available within the forest floor from mineralization of organic matter. Virtually all the available ammonium (mineralized plus input from throughfall) is utilized in the forest floor, either by microorganisms or through uptake by vegetation. The most abundant N component of soil water solutions leaving the system was nitrate. Net ecosystem fluxes indicate accumulation of both ammonium and nitrate. There is a small net loss of organic N from the ecosystem. Some nitrate leaves the bottom of the B horizon throughout the year. Comparisons with other temperate coniferous sites and examination of the ecosystem N mass balance indicate that N use efficiency is less at our site, which suggests that the site is not severely limited by N.     

Nitrogen cycling and anthropogenic impact in the tropical interamerican seas

We discuss the mechanisms leading to nutrient limitation in tropical marine systems, with particular emphasis on nitrogen cycling in Caribbean ecosystems. We then explore how accelerated nutrient cycling from human activities is affecting these systems.Both nitrogen and phosphorus exert substantial influence on biological productivity and structure of tropical marine ecosystems. Offshore planktonic communities are largely nitrogen limited while nearshore ecosystems are largely phosphorus limited. For phosphorus, the ability of sediment to adsorb and store phosphorus is probably greater for tropical carbonate sediments than for most nearshore sediments in temperate coastal systems. However, the ability of tropical carbonate sediments to take up phosphorus can become saturated as phosphorus loading from human sources increases. The nature of the sediment, the mixing rate between nutrient-laden runoff waters and nutrient-poor oceanic waters and the degree of interaction of these water masses with the sediment will probably control the dynamics of this transition.Nearshore tropical marine ecosystems function differently from their temperate counterparts where coupled nitrification/denitrification serves as an important mechanism for nitrogen depuration. In contrast, nearshore tropical ecosystems are more susceptible to nitrogen loading as depurative capacity of the microbial communities is limited by the fragility of the nitrification link. At the same time, accumulation of organic matter in nearshore carbonate sediments appears to impair their capacity for phosphorus immobilization. In the absence of depurative mechanisms for either phosphorus or nitrogen, limitation for both these nutrients is alleviated and continued nutrient loading fuels the proliferation of nuisance algae.     

Fate and Transport of Organic Nitrogen in Minimally Disturbed Montane Streams of Colorado,USA

In two montane watersheds that receive minimal deposition of atmospheric nitrogen, 15–71% of dissolved organic nitrogen (DON) was bioavailable in stream water over a 2-year period. Discharge-weighted concentrations of bulk DON were between 102 and 135 μg/l, and the C:N ratio differed substantially between humic and non-humic fractions of DON. Approximately 70% of DON export occurred during snowmelt, and 40% of that DON was biologically available to microbes in stream sediments. Concentrations of bioavailable DON in stream water were 2–16 times greater than dissolved inorganic nitrogen (DIN) during the growing season, and bioavailable DON was depleted within 2–14 days during experimental incubations. Uptake of DON was influenced by the concentration of inorganic N in stream water, the concentration of non-humic DON in stream water, and the C:N ratio of the non-humic fraction of dissolved organic matter (DOM). Uptake of DON declined logarithmically as the concentration of inorganic N in stream water increased. Experimental additions of inorganic N also caused a decline in uptake of DON and net production of DON when the C:N ratio of non-humic DOM was high. This study indicates that the relative and absolute amount of bioavailable DON can vary greatly within and across years due to interactions between the availability of inorganic nutrients and composition of DOM. DOM has the potential to be used biotically at a high rate in nitrogen-poor streams, and it may be generated by heterotrophic microbes when DIN and labile DOM with low relative nitrogen content become abundant.     

Grazing history effects on above- and below-ground litter decomposition and nutrient cycling in two co-occurring grasses

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment.     

Nitrogen cycling in Louisiana Gulf Coast brackish marshes

Nitrogen fixation and nitrogen accumulation were measured in a Louisiana Spartina patens brackish marsh. Using the acetylene reduction technique calibrated with direct ¹⁵N₂ assimilation, an equivalent of 90.0 µ g N g^–1 yr^–1 was fixed. Fixation was greater in the summer months and in the upper portion of the soil profile. Extractable ammonium increased with depth and was negatively correlated with ethylene production. Average ammonium concentration in the sediment was 39 µg NH₄ ⁺-N g^–1 sediment. Cesium-137 dating of the soil profile showed the marsh was vertically accreting at a rate of 0.60 cm yr^–1. Calculations using vertical accretion rate, bulk density, and total nitrogen content of sediment indicate that the marshes are accumumating 7.2 g Nm^–2 yr^–1 thus serving as a major nitrogen sink. Measured nitrogen fluxes were incorporated with existing flux measurement in developing a nitrogen budget for the marsh.     

"稻鸭共生"生态系统稻季N、P循环

稻鸭共生是对我国传统农业稻田养鸭的继承与发展。在长江流域双季稻主产区湖南布置了稻田养鸭田间试验,以常规稻作为对照,研究"稻鸭共生"生态系统N、P循环。结果表明:"稻鸭共生"生态系统N、P输出分别为153.50 kg/hm²和59.03 kg/hm²,其中鸭产品N、P输出分别是23.98 kg/hm²和3.10 kg/hm²;"稻鸭共生"系统在目前的N、P养分投入水平下,土壤存在严重的N和P亏缺;鸭子系统N、P输入对系统外饲料投入依赖高,自持能力较差;鸭粪N、P参与当季的稻田养分循环,其循环率分别为10.66%和28.16%。     

Nitrogen cycling in grazed pastures at elevated CO2: N returns by ruminants

In pastures grazed by large herbivores, nutrients cycle both through litter and animal excreta. We compared nitrogen (N) returns from sheep grazing a temperate pasture exposed to ambient or elevated CO₂ (475 μmol mol^?1) in a FACE (Free Air CO₂ Enrichment) experiment established in the spring of 1997. In the spring of 2000 and 2001, we measured the chemical composition of the diet, sheep faeces and of individual plant species before grazing to characterize feed intake and to compare the intake of N to the N produced in faeces. In both years under elevated CO₂, leaves of the individual species exhibited lower N concentrations and higher water‐soluble carbohydrate (WSC) concentrations. There was a significantly greater proportion of legume in the diet at elevated CO₂ but, together with the changes in chemical composition of individual species, this resulted in diets that had similar N but higher WSC and digestibility for both ambient and elevated CO₂. We found that a greater proportion of dietary N was partitioned to urine at elevated CO₂, probably because of the higher proportion of legume N in the diet, with possible differences in protein quality. A potentially significant consequence of this change in partitioning is greater N loss through volatilization at higher CO₂ levels.     

Ammonia volatilization and the effects of large grazing mammals on nutrient loss from East African grasslands

Summary Ammonia volatilization losses measured from soils at seven sites in the Serengeti National Park, Tanzania during the 1986 growing season ranged from 2.78±0.49% to 25.03±1.34% of nitrogen applied. Although peak ammonia losses ranged from 0.071±0.018 to 0.404±0.040 g N m^-2 h^-1, rates dropped to zero within four days, and calculations reveal that volatilization losses represent minor fluxes in the context of the system's nitrogen cycling. Volatilization losses were inversely correlated with grazing intensity experienced by a site, and it appears that large ungulates themselves contribute to nutrient conservation throught indirect interactive effects on system processes.