Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.     

Does nitrogen addition to raised bogs influence peat phosphorus pools?

Two Sphagnum moss species occupying hummock areas (Sphagnum capillifolium) and wetter hollows (Sphagnum recurvum) on a raised bog in north east Scotland were treated every two weeks with NH₄NO₃ solutions to supply 3g N m^–2 yr^–1. Although S. recurvum moss contained a greater concentration of total P than S. capillifolium the amounts and N:P ratios were similar in both species. Larger amounts of total dissolved P (TDP) and molybdate reactive P (MRP) were extracted from beneath S. recurvum to 25 cm below the moss. Additions of N both increased and decreased the amounts of TDP at different times, and decreased MRP. The MRP fraction accounted for about 20 per cent of TDP and the difference was assumed to be in organic forms (DOP). Nitrogen addition had no effect on the amounts of DOP, but C:P ratios of this fraction changed with species, depth and N addition. Microbial P accounted for as much as 70 per cent of total P and showed seasonal variations, but no differences between the two moss species and N addition.     

Vertical transport of dissolved organic C and N under long-term N amendments in pine and hardwood forests

At the Harvard Forest, Massachusetts, a long-term effort is under way to study responses in ecosystem biogeochemistry to chronic inputs of N in atmospheric deposition in the region. Since 1988, experimental additions of NH₄NO₃ (0, 5 and 15 g N m^–2 yr^–1) have been made in two forest stands:Pinus resinosa (red pine) and mixed hardwood. In the seventh year of the study, we measured solute concentrations and estimated solute fluxes in throughfall and at two soil depths, beneath the forest floors (Oa) and beneath the B horizons.Beneath the Oa, concentrations and fluxes of dissolved organic C and N (DOC and DON) were higher in the coniferous stand than in the hardwood stand. The mineral soil exerted a strong homogenizing effect on concentrations beneath the B horizons. In reference plots (no N additions), DON composed 56% (pine) and 67% (hardwood) of the total dissolved nitrogen (TDN) transported downward from the forest floor to the mineral soil, and 98% of the TDN exported from the solums. Under N amendments, fluxes of DON from the forest floor correlated positively with rates of N addition, but fluxes of inorganic N from the Oa exceeded those of DON. Export of DON from the solums appeared unaffected by 7 years of N amendments, but as in the Oa, DON composed smaller fractions of TDN exports under N amendments. DOC fluxes were not strongly related to N amendment rates, but ratios of DOC:DON often decreased.The hardwood forest floor exhibited a much stronger sink for inorganic N than did the pine forest floor, making the inputs of dissolved N to mineral soil much greater in the pine stand. Under the high-N treatment, exports of inorganic N from the solum of the pine stand were increased >500-fold over reference (5.2 vs. 0.01 g N m^–2 yr^–1), consistent with other manifestations of nitrogen saturation. Exports of N from the solum in the pine forest decreased in the order NO₃-N> NH₄-N> DON, with exports of inorganic N 14-fold higher than exports of DON. In the hardwood forest, in contrast, increased sinks for inorganic N under N amendments resulted in exports of inorganic N that remained lower than DON exports in N-amended plots as well as the reference plot.     

Clinoptilolite zeolite and cellulose amendments to reduce ammonia volatilization in a calcareous sandy soil

Leaching of nitrate (NO₃ ^–) below the root zone and gaseous losses of nitrogen (N) such as ammonia (NH₃) volatilization, are major mechanisms of N loss from agricultural soils. New techniques to minimize such losses are needed to maximize N uptake efficiency and minimize production costs and the risk of potential N contamination of ground and surface waters. The effects of cellulose (C), clinoptilolite zeolite (CZ), or a combination of both (C+CZ) on NH₃ volatilization and N transformation in a calcareous Riviera fine sand (loamy, siliceous, hyperthermic, Arenic Glossaqualf) from a citrus grove were investigated in a laboratory incubation study. Ammonia volatilization from NH₄NO₃ (AN), (NH₄)₂SO₄(AS), and urea (U) applied at 200 mg N kg^–1 soil decreased by 2.5-, 2.1- and 0.9-fold, respectively, with cellulose application at 15 g kg^–1 and by 4.4-, 2.9- and 3.0-fold, respectively, with CZ application at 15 g kg^–1 as compared with that from the respective sources without the amendments. Application of cellulose plus CZ (each at 15 g kg^–1) was the most effective in decreasing NH₃ volatilization. Application of cellulose increased the microbial biomass, which was responsible for immobilization of N, and thus decreased volatilization loss of NH₃–N. The effect of CZ, on the other hand, may be due to increased retention of NH₄ in the ion-exchange sites. The positive effect of interaction between cellulose and CZ amendment on microbial biomass was probably due to improved nutrient retention and availability to microorganisms in the soil. Thus, the amendments provide favorable conditions for microbial growth. These results indicate that soil amendment of CZ or CZ plus organic materials such as cellulose has great potential in reducing fertilizer N loss in sandy soils.     

Ammonium,nitrate and dissolved organic nitrogen in seepage water as affected by compost amendment to European beech,Norway spruce,and Scots pine forests

Fertilization of nutrient-depleted and degraded forest soils may be required to sustain utilization of forests. In some European countries, the application of composts may now be an alternative to the application of inorganic fertilizers because commercial compost production has increased and compost quality has been improved. There is, however, concern that compost amendments may cause increased leaching of nitrogen, trace metals and toxic organic compounds to groundwater. The objective of this study was to assess the risk of ammonium (NH₄ ⁺), nitrate (NO₃ ^–) and dissolved organic nitrogen (DON) leaching following a single compost application to silty and sandy soils in mature beech (Fagus sylvatica L.), pine (Pinus silvestris L.) and spruce (Picea abies Karst.) forests at Solling and Unterlüß in Lower Saxony, Germany. Mature compost from separately collected organic household waste was applied to the soil surface at a rate of 6.3 kg m^–2 in the summer of 1997 and changes in NH₄ ⁺, NO₃ ^– and DON concentrations in throughfall and soil water at 10 and 100 cm soil depths were determined for 32 months. The spruce forests had the highest N inputs by throughfall water and the highest N outputs in both the control and compost plots compared with the pine and beech forests. Overall, the differences in total N outputs at 100 cm soil depth between the control and compost plots ranged between 0.3 and 11.2 g N m^–2 for the entire 32-month period. The major leaching of these amounts occurred during the first 17 months after compost amendments, but there was no significant difference in total N outputs (–0.2 to 1.8 g N m^–2) between the control and compost plots during the remaining 15 months. Most of the mineral soils acted as a significant sink for NO₃ ^– and DON as shown by a reduction of their outputs from 10 to 100 cm depth. Based on these results, we conclude that application of mature compost with high inorganic N contents could diminish the groundwater quality in the first months after the amendments. A partial, moderate application of mature compost with low inorganic N content to nutrient depleted forest soils can minimize the risk of NO₃ ^– leaching.     

The Role of Dissolved Organic Carbon, Dissolved Organic Nitrogen, and Dissolved Inorganic Nitrogen in a Tropical Wet Forest Ecosystem

Although tropical wet forests play an important role in the global carbon (C) and nitrogen (N) cycles, little is known about the origin, composition, and fate of dissolved organic C (DOC) and N (DON) in these ecosystems. We quantified and characterized fluxes of DOC, DON, and dissolved inorganic N (DIN) in throughfall, litter leachate, and soil solution of an old-growth tropical wet forest to assess their contribution to C stabilization (DOC) and to N export (DON and DIN) from this ecosystem. We found that the forest canopy was a major source of DOC (232 kg C ha^–1 y^–1). Dissolved organic C fluxes decreased with soil depth from 277 kg C ha^–1 y^–1 below the litter layer to around 50 kg C kg C ha^–1 y^–1 between 0.75 and 3.5m depth. Laboratory experiments to quantify biodegradable DOC and DON and to estimate the DOC sorption capacity of the soil, combined with chemical analyses of DOC, revealed that sorption was the dominant process controlling the observed DOC profiles in the soil. This sorption of DOC by the soil matrix has probably led to large soil organic C stores, especially below the rooting zone. Dissolved N fluxes in all strata were dominated by mineral N (mainly NO₃⁻). The dominance of NO₃^– relative to the total amount nitrate of N leaching from the soil shows that NO₃^– is dominant not only in forest ecosystems receiving large anthropogenic nitrogen inputs but also in this old-growth forest ecosystem, which is not N-limited.     

Dissolved organic carbon affects soil microbial activity and nitrogen dynamics in a Mexican tropical deciduous forest

Seasonal variation of dissolved organic C (DOC) and its effects on microbial activity and N dynamics were studied during two consecutive years in soils with different organic C concentrations (hilltop and hillslope) in a tropical deciduous forest of Mexico. We found that DOC concentrations were higher at the hilltop than at the hillslope soils, and in both soils generally decreased from the dry to the rainy season during the two study years. Microbial biomass and potential C mineralization rates, as well as dissolved organic N (DON) and NH₄⁺ concentrations and net N immobilization were higher in soils with higher DOC than in soils with lower DOC. In contrast, net N immobilization and NH₄⁺ concentration were depleted in the soil with lowest DOC, whereas NO₃⁻ concentrations and net nitrification increased. Negative correlations between net nitrification and DOC concentration suggested that NH₄⁺ was transformed to NO₃⁻ by nitrifiers when the C availability was depleted. Taken together, our results suggest that available C appears to control soil microbial activity and N dynamics, and that microbial N immobilization is facilitated by active heterotrophic microorganisms stimulated by high C availability. Soil autotrophic nitrification is magnified by decreases in C availability for heterotrophic microbial activity. This study provides an experimental data set that supports the conceptual model to show and highlight that microbial dynamics and N transformations could be functionally coupled with DOC availability in the tropical deciduous forest soils. Responsible Editor: Chris Neill     

Effects of forest clear-cutting on the carbon and nitrogen fluxes through podzolic soil horizons

Effects of clear-cutting on the dissolved fluxes of organic C (DOC), organic N (DON), NO₃ ^– and NH₄ ⁺ through surface soil horizons were studied in a Norway spruce dominated mixed boreal forest in eastern Finland. Bulk deposition, total throughfall and soil water from below the organic (including understorey vegetation and, after clear-cutting, also logging residues), eluvial and illuvial horizons were sampled weekly from 1993 to 1999. Clear-cutting was carried out in September 1996. The removal of the tree canopy decreased the deposition of DOC and DON to the forest floor and increased that of NH₄ ⁺ and NO₃ ^– but did not affect the deposition of total N (DTN, <3 kg ha^–1 a^–1). The leaching of DOC and DON from the organic horizon increased over twofold after clear-cutting (fluxes were on an average 168 kg C and 3.3 kg N ha^–1 a^–1), but the increased outputs were effectively retained in the surface mineral soil horizons. Inorganic N deposition was mainly retained by the logging residues and organic horizon indicating microbial immobilization. Increased NO₃ ^– formation reflected as elevated concentrations in the percolate from below the mineral soil horizons were observed especially in the third year after clear-cutting. However, the changes were small and the increased leaching of DTN from below the illuvial horizon remained small (<0.4 kg ha^–1 a^–1) and mainly DON. Effects of clear-cutting on the transport of C and N to surface waters will probably be negligible.     

Growth and solute composition of the salt-tolerant kallar grass [Leptochloa fusca (L.) Kunth] as affected by nitrogen source

A study was conducted to elucidate the effect of N form, either NH₄ ⁺ or NO₃ ^–, on growth and solute composition of the salt-tolerant kallar grass [Leptochloa fusca (L.) Kunth] grown under 10 mM or 100 mM NaCl in hydroponics. Shoot biomass was not affected by N form, whereas NH₄ ⁺ compared to NO₃ ^– nutrition caused an almost 4-fold reduction in the root biomass at both salinity levels. Under NH₄ ⁺ nutrition, salinity had no effect on the biomass yield, whereas under NO₃ ^– nutrition, increasing salinity from 10 mM to 100 mM caused 23% and 36% reduction in the root and shoot biomass, respectively. The reduced root growth under NH₄ ⁺ nutrition was not attributable to impaired shoot to root C allocation since N form did not affect the overall root sugar concentration and the starch concentration was even higher under NH₄ ⁺ compared to NO₃ ^– nutrition. The low NH₄ ⁺ (2 mM) and generally higher amino-N concentrations in NH₄ ⁺- compared to NO₃ ^–-fed plants indicated that the grass was able to effectively detoxify NH₄ ⁺. Salinity had no effect on Ca²⁺ and Mg²⁺ levels, whereas their concentration in shoots was lower under NH₄ ⁺ compared to NO₃ ^– nutrition (over 66% reduction in Ca²⁺; over 20% reduction in Mg²⁺), but without showing deficiency symptoms. Ammonium compared to NO₃ ^– nutrition did not inhibit K⁺ uptake, and the K⁺-Na⁺ selectivity either remained unaffected or it was higher under NH₄ ⁺ than under NO₃ ^– nutrition. Results suggested that while NH₄ ⁺ versus NO₃ ^– nutrition substantially reduced root growth, and also strongly modified anion concentrations and to a minor extent concentrations of divalent cations in shoots, it did not influence salt tolerance of kallar grass.     

Microbial activity under alpine snowpacks, Niwot Ridge, Colorado

Experiments were conducted during 1993 at Niwot Ridge in the Colorado Front Range to determine if the insulating effect of winter snow cover allows soil microbial activity to significantly affect nitrogen inputs and outputs in alpine systems. Soil surface temperatures under seasonal snowpacks warmed from –14 °C in January to 0 °C by May 4th. Snowmelt began in mid-May and the sites were snow free by mid June. Heterotrophic microbial activity in snow-covered soils, measured as C0₂ production, was first identified on March 4, 1993. Net C0₂ flux increased from 55 mg CO₂-C m^–2 day^–1 in early March to greater than 824 mg CO₂-C m-2 day^–1 by the middle of May. Carbon dioxide production decreased in late May as soils became saturated during snowmelt. Soil inorganic N concentrations increased before snowmelt, peaking between 101 and 276 mg kg^–1 soil in May, and then decreasing as soils became saturated with melt water. Net N mineralization for the period of March 3 to May 4 ranged from 2.23 to 6.63 g N m^–2, and were approximately two orders of magnitude greater than snowmelt inputs of 50.4 mg N m^–2 for NH₄ ⁺ and 97.2 mg N m^–2 for NO₃ ^–. Both NO₃ ^– and NH₄ ⁺ concentrations remained at or below detection limits in surface water during snowmelt, indicating the only export of inorganic N from the system was through gaseous losses. Nitrous oxide production under snow was first observed in early April. Production increased as soils warned, peaking at 75 g N₂O-N m^–2 day^–1 in soils saturated with melt water one week before the sites were snow free. These data suggest that microbial activity in snow-covered soils may play a key role in alpine N cycling before plants become active.     

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.     

Carbon and nitrogen cycling during old-field succession: Constraints on plant and microbial biomass

Soil C and N dynamics were studied in a sequence of old fields of increasing age to determine how these biogeochemical cycles change during secondary succession. In addition, three different late-successional forests were studied to represent possible "steady state" conditions. Surface soil samples collected from the fields and forests were analyzed for total C, H₂O-soluble C, total N, potential net N mineralization, potential net nitrification, and microbial biomass. Above-and belowground plant biomass was estimated within each of the old field sites.Temporal changes in soil organic C, total N and total plant biomass were best described by a gamma function [y =at ^b e ^ctd +f] whereas a simple exponential model [y =a(l – e^–bt ) + c] provided the best fit to changes in H₂O-soluble C, C:N ratio, microbial C, and microbial N. Potential N mineralization and nitrification linearly increased with field age; however, rates were variable among the fields. Microbial biomass was highly correlated to soil C and N pools and well correlated to the standing crop of plant biomass. In turn, plant biomass was highly correlated to pools and rates of N cycling.Patterns of C and N cycling within the old field sites were different from those in a northern hardwood forest and a xeric oak forest; however, nutrient dynamics within an oak savanna were similar to those found in a 60-yr old field. Results suggest that patterns in C and N cycling within the old-field chronosequence were predictable and highly correlated to the accrual of plant and microbial biomass.     

Landscape Controls on Organic and Inorganic Nitrogen Leaching across an Alpine/Subalpine Ecotone,Green Lakes Valley,Colorado Front Range

Here we report measurements of organic and inorganic nitrogen (N) fluxes from the high-elevation Green Lakes Valley catchment in the Colorado Front Range for two snowmelt seasons (1998 and 1999). Surface water and soil samples were collected along an elevational gradient extending from the lightly vegetated alpine to the forested subalpine to assess how changes in land cover and basin area affect yields and concentrations of ammonium-N (NH₄-N), nitrate-N (NO₃-N), dissolved organic N (DON), and particulate organic N (PON). Streamwater yields of NO₃-N decreased downstream from 4.3 kg ha⁻¹ in the alpine to 0.75 kg ha⁻¹ at treeline, while yields of DON were much less variable (0.40–0.34 kg ha⁻¹). Yields of NH₄-N and PON were low and showed little variation with basin area. NO₃-N accounted for 40%–90% of total N along the sample transect and was the dominant form of N at all but the lowest elevation site. Concentrations of DON ranged from approximately 10% of total N in the alpine to 45% in the subalpine. For all sites, volume-weighted mean concentrations of total dissolved nitrogen (TDN) were significantly related to the DIN:DON ratio (R ² = 0.81, P < 0.001) Concentrations of NO₃-N were significantly higher at forested sites that received streamflow from the lightly vegetated alpine reaches of the catchment than in a control catchment that was entirely subalpine forest, suggesting that the alpine may subsidize downstream forested systems with inorganic N. KCl-extractable inorganic N and microbial biomass N showed no relationship to changes in soil properties and vegetative cover moving downstream in catchment. In contrast, soil carbon–nitrogen (C:N) ratios increased with increasing vegetative cover in catchment and were significantly higher in the subalpine compared to the alpine (P < 0.0001) Soil C:N ratios along the sample transect explained 78% of the variation in dissolved organic carbon (DOC) concentrations and 70% of the variation in DON concentrations. These findings suggest that DON is an important vector for N loss in high-elevation ecosystems and that streamwater losses of DON are at least partially dependent on catchment soil organic matter stoichiometry. Received 26 July 2001; accepted 6 May 2002.     

Controls on the dynamics of dissolved organic carbon and nitrogen in a Central European deciduous forest

Despite growing attention concerning therole of dissolved organic matter (DOM) inelement cycling of forest ecosystems, thecontrols of concentrations and fluxes of bothdissolved organic carbon (DOC) and nitrogen(DON) under field conditions in forest soilsremain only poorly understood. The goal ofthis project is to measure the concentrations and fluxes of DON, NH₄ ⁺, NO₃ ^–and DOC in bulkprecipitation, throughfall, forest floorleachates and soil solutions of a deciduousstand in the Steigerwald region (northernBavaria, Germany). The DOC and DONconcentrations and fluxes were highest inleachates originating from the Oa layer of theforest floor (73 mg C L^–1, 2.3 mg NL^–1 and about 200–350 kg C, 8–10 kg Nha^–1 yr^–1). They were observed to behighly variable over time and decreased in themineral topsoil (17 mg C L^–1, 0.6 mg NL^–1 and about 50–90 kg C, 2.0 to 2.4 kg Nha^–1 yr^–1). The annual variability ofDOC and DON concentrations and subsequentialDOC/DON ratios was substantial in allsolutions. The DOC and DON concentrations inthroughfall were positively correlated withtemperature. The DOC and DON concentrationsdid not show seasonality in the forest floorand mineral soil. Concentrations were notrelated to litterfall dynamics but didcorrespond in part to the input of DOC and DONfrom throughfall. The throughfall contributionto the overall element fluxes was higher forDON than for DOC. Concentrations and fluxes ofDON were significantly correlated to DOC inthroughfall and the Oi layer. However, thecorrelation was weak in Oa leachates. Inaddition, seasonal and annual variation ofDOC/DON ratios indicated different mechanismsand release rates from the forest floor forboth components. The concentrations of DOC andDON in forest floor leachates were in mostcases dependent neither on the pH value orionic strength of the solution, nor on thewater flux or temperature changes. As aconsequence, the DOC and DON fluxes from theforest floor into the mineral soil werelargely dependent on the water flux if annualand biweekly time scales are considered.     

A comparison of organic and inorganic nitrogen fertilizers: Their nitrate-N and ammonium-N release characteristics and effects on the growth response of lettuce (Lactuca sativa L. cv. Fortune)

The response of pot grown lettuce to inorganic (ammonium nitrate) and organic (dried blood and Protox) N fertilizers was determined at two temperature regimes (15°C day/10°C night and 20°C day/15°C night) and related to the NH₄–N and NO₃–N release characteristics of each material. The N release characteristics of the organic materials matched the N requirements of lettuce more closely than the inorganic fertilizer. However, was rapidly released from the protein based materials such that composts were depleted of available fertilizer N at the same time irrespective of the form supplied. The warmer temperature regimes resulted in a more rapid depletion of the fertilizers due to biological immobilization such that N recoveries in shoots, roots and leachates were reduced. Approximately 20% of the N present in Protox (a material derived from activated sewage sludge, processed to reduce the heavy metal content to minimal levels) appeared to be resistant to microbial degradation and was unavailable to the plants. Therefore, the growth response of lettuce was slightly reduced with Protox compared to the other materials at similar rates of incorporation. The organic materials did not contribute NO₃–N to the plant and small NO₃–N concentrations in petioles were derived from the water used for irrigation. However, NO₃–N levels in plants receiving inorganic ammonium nitrate were initially high but progressively declined as the fertilizer NO₃–N became depleted.     

Recruitment preferences of blue mussel spat (Mytilus edulis) for different substrata and microhabitats in the White Sea (Russia)

We adapted the chloroform fumigation method to determine microbial nitrogen (N) and microbial incorporation of ¹⁵N on three common substrates [leaves, wood and fine benthic organic matter (FBOM)] in three forest streams. We compared microbial N and ¹⁵N content of samples collected during a 6-week ¹⁵N–NH₄ tracer addition in each stream. The ¹⁵N was added during late autumn to Upper Ball Creek, a second-order stream at the Coweeta Hydrologic Lab, North Carolina, U.S.A.; during spring to Walker Branch, a first-order stream on DOE's Oak Ridge National Environmental Research Park, Tennessee; and during summer to Bear Brook, a first-order stream in the Hubbard Brook Experimental Forest, New Hampshire. FBOM was the largest component of organic matter and N standing stock in all streams. Microbial N represented the highest proportion of total N in leaves and least in FBOM in Walker Branch and Bear Brook. In Upper Ball Creek, the proportion of microbial N was higher in FBOM than in used biofilm or on leaves. Standing stock of microbial N on leaves and in FBOM ranged from 37 mg N m^–2 in Bear Brook to 301 mg N m^–2 in Walker Branch. Percent of detrital N in living microbial cells was directly related to total microbial biomass (fungal and bacterial biomass) determined from microscopic counts. ¹⁵N values for microbes were generally higher than for bulk detritus, which would result in higher ¹⁵N values for animals preferentially consuming or assimilating microbial cells. The proportion of ¹⁵N taken up by detritus during the ¹⁵N experiments that remained in microbial cells by the end of the experiments was highest for wood biofilm in Upper Ball Creek (69%), leaves in Walker Branch (65%) and FBOM in Upper Ball Creek (31%). Lower retention proportions (<1–25%) were observed for other substrates. Our results suggest that microbial cells associated with leaves and wood biofilm were most active in ¹⁵N–NH₄ immobilization, whereas microbial cells associated with FBOM immobilized little ¹⁵N from stream water.     

Patterns of restoration of soil physciochemical properties and microbial biomass in different landslide sites in the sal forest ecosystem of Nepal Himalaya

The pattern of natural restoration in soil components and processes was documented in five landslide-damaged (1–58-year-old) sites in the moist tropical sal (Shorea robusta) forest ecosystem of Nepal Himalaya. Comparisons were made with an undisturbed forest site in the same region. Concentrations of soil organic C, total N, total P and extractable nutrients (Ca, Mg and K) increased with the age of sites. The 58-year-old site showed concentrations of soil organic C, total N and total P that were 75–89% of concentrations in the undisturbed sal forest. The soil microbial biomass, the active fraction of soil organic matter, showed similar seasonal variations at all sites. The amount of mean microbial biomass (expressed as C, N and P contents) increased 4–5 times at the 58-year-old site relative to the 1-year-old site, and the bulk increase occurred within the initial 15 year. The increase in the C/N ratio of soil microbial biomass with age (9.4–11.6 years) reflected change in its composition. Although the net N-mineralization rate increased consistently until 58 years of age, the proportion of nitrification rate relative to ammonification rate distinctly decreased beyond 40 years. On the other hand, the soil available-N (both NO₃⁻ and NH₄⁺) concentrations increased from 1 to 40 year and then declined; with age the proportion of NH₄⁺ increased, however. Rates of restoration in soil properties were faster in the early successional stages (1–15 year) than late stages. Among different soil properties the restoration of soil microbial biomass (C and N) was faster than soil organic C and total N. Best fit power function models showed that the estimated times for the 58-year-old site to reach the level of the undisturbed, mature sal forest would be about 30–35 year for microbial biomass (C and N) and about 100–150 year for organic C and total N. Higher accumulation of soil microbial biomass and high N-mineralization rate at late successional stages indicated the re-establishment of enriched soil and restitution of nutrient cycling during the course of ecosystem restoration.     

Nitrification in Dutch heathland soils

A survey was conducted over a range of 17 Dutch heathland locations, subdivided into 41 sites dominated by either dwarf-shrubs (Calluna vulgaris or Erica tetralix) or grass species (Deschampsia flexuosa or Molinia caerulea). Among the habitats of the dominant plant species relatively little differences in general soil properties were observed. The P status of Deschampsia sites was relatively high as well as the NO₃ ^–-N concentrations in the 0–10 cm layer (FH included) at the grass-dominated sites. At sites with a dead or degenerating dwarf-shrub vegetation, NH₄ ⁺-N concentrations reached very high levels.Net production of nitrate was observed during incubation of intact 0–10 cm soil cores (FH-layer included) in the laboratory for all sites, even though in some instances, particularly at Calluna and Erica sites, no nitrate was initially measured. Generally, a higher nitrification rate was found for the grass-dominated sites, and for Deschampsia in particular. The net production of nitrate was highly significantly correlated with net N mineralization, being a reasonable predictor of nitrification in a simple regression model (R²=0.47; P<0.001). Net nitrification was also significantly correlated with the NO₃ ^–-N initially present at the start of the growing season (R=0.65; P<0.001) and with the labile organic P content of the soil (R=0.65; P<0.001). By including initial NO₃ ^–-N and labile organic P, together with net N mineralization and pH, in a multiple regression model, net nitrate production could be predicted with a much higher precision (R²=0.75; P<0.001). Although apparent nitrification was not significantly correlated with pH, the latter contributed significantly to the multiple regression equation for the prediction of the former.The influence of the labile organic P pool may act via its positive correlation with microbial biomass, thus more or less reflecting the potential mineralization/nitrifying capacity of a particular site.     

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     

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.