Inputs,losses and transformations of nitrogen and phosphorus in the pelagic North Atlantic Ocean

The North Atlantic Ocean receives the largest allochthonous supplies of nitrogen of any ocean basin because of the close proximity of industrialized nations. In this paper, we describe the major standing stocks, fluxes and transformations of nitrogen (N) and phosphorus (P) in the pelagic regions of the North Atlantic, as one part of a larger effort to understand the entire N and P budgets in the North Atlantic Ocean, its watersheds and overlying atmosphere. The primary focus is on nitrogen, however, we consider both nitrogen and phosphorus because of the close inter-relationship between the N and P cycles in the ocean. The oceanic standing stocks of N and P are orders of magnitude larger than the annual amount transported off continents or deposited from the atmosphere. Atmospheric deposition can have an impact on oceanic nitrogen cycling at locations near the coasts where atmospheric sources are large, or in the centers of the highly stratified gyres where little nitrate is supplied to the surface by vertical mixing of the ocean. All of the reactive nitrogen transported to the coasts in rivers is denitrified or buried in the estuaries or on the continental shelves and an oceanic source of nitrate of 0.7–0.95 × 10¹² moles NO ₃ ^–1 y^–1 is required to supply the remainder of the shelf denitrification (Nixon et al., this volume). The horizontal fluxes of nitrate caused by the ocean circulation are both large and uncertain. Even the sign of the transport across the equator is uncertain and this precludes a conclusion on whether the North Atlantic Ocean as a whole is a net source or sink of nitrate. We identify a source of nitrate of 3.7–6.4 × 10¹² moles NO ₃ ^– y^–1 within the main thermocline of the Sargasso Sea that we infer is caused by nitrogen fixation. This nitrate source may explain the nitrate divergence observed by Rintoul & Wunsch (1991) in the mid-latitude gyre. The magnitude of nitrogen fixation inferred from this nitrate source would exceed previous estimates of global nitrogen fixation. Nitrogen fixation requires substantial quantities of iron as a micro-nutrient and the calculated iron requirement is comparable to the rates supplied by the deposition of iron associated with Saharan dust. Interannual variability in dust inputs is large and could cause comparable signals in the nitrogen fixation rate. The balance of the fluxes across the basin boundaries suggest that the total stocks of nitrate and phosphate in the North Atlantic may be increasing on time-scales of centuries. Some of the imbalance is related to the inferred nitrogen fixation in the gyre and the atmospheric deposition of nitrogen, both of which may be influenced by human activities. However, the fluxes of dissolved organic nutrients are almost completely unknown and they have the potential to alter our perception of the overall mass balance of the North Atlantic Ocean.     

Estimating denitrification in North Atlantic continental shelf sediments

A model of coupled nitrification/denitrification was developed for continental shelf sediments to estimate the spatial distribution of denitrification throughout shelf regions in the North Atlantic basin. Using data from a wide range of continental shelf regions, we found a linear relationship between denitrification and sediment oxygen uptake. This relationship was applied to specific continental shelf regions by combining it with a second regression relating sediment oxygen uptake to primary production in the overlying water. The combined equation was: denitrification (mmol N m^–2 d^–1)=0.019^* phytoplankton production (mmol C m^–2 d^–1). This relationship suggests that approximately 13% of the N incorporated into phytoplankton in shelf waters is eventually denitrified in the sediments via coupled nitrification/denitrification, assuming a C:N ratio of 6.625:1 for phytoplankton. The model calculated denitrification rates compare favorably with rates reported for several shelf regions in the North Atlantic.The model-predicted average denitrification rate for continental shelf sediments in the North Atlantic Basin is 0.69 mmol N m^{– 2} d^–1. Denitrification rates (per unit area) predicted by the model are highest for the continental shelf region in the western North Atlantic between Cape Hatteras and South Florida and lowest for Hudson Bay, the Baffin Island region, and Greenland. Within latitudinal belts, average denitrification rates were lowest in the high latitudes, intermediate in the tropics and highest in the mid-latitudes. Although denitrification rates per unit area are lowest in the high latitudes, the total N removal by denitrification (53 × 10¹⁰ mol N y^–1) is similar to that in the mid-latitudes (60 × 10¹⁰ mol N y^–1) due to the large area of continental shelf in the high latitudes. The Gulf of St. Lawrence/Grand Banks area and the North Sea are responsible for seventy-five percent of the denitrification in the high latitude region. N removal by denitrification in the western North Atlantic (96 × 10¹⁰ mol N y^–1) is two times greater than in the eastern North Atlantic (47 × 10¹⁰ mol N y^–1). This is primarily due to differences in the area of continental shelf in the two regions, as the average denitrification rate per unit area is similar in the western and eastern North Atlantic.We calculate that a total of 143 × 10¹⁰ mol N y^–1 is removed via coupled nitrification/denitrification on the North Atlantic continental shelf. This estimate is expected to underestimate total sediment denitrification because it does not include direct denitrification of nitrate from the overlying water. The rate of coupled nitrification/denitrification calculated is greater than the nitrogen inputs from atmospheric deposition and river sources combined, and suggests that onwelling of nutrient rich slope water is a major source of N for denitrification in shelf regions. For the two regions where N inputs to a shelf region from onwelling have been measured, onwelling appears to be able to balance the denitrification loss.     

Atmospheric deposition of nutrients to the North Atlantic Basin

Atmospheric chemical models are used to estimate the deposition rate of various inorganic oxides of nitrogen (NO_y), reduced nitrogen species (NH_x) and mineral dust to the North Atlantic Ocean (NAO). The estimated deposition of NO_y to the NAO (excluding the coastal ocean) and the Caribbean is 360 × 10⁹ Moles-N m^–2 yr^–1 (5.0 Tg N); this is equivalent to about 13% of the estimated global emission rate (natural and anthropogenic) and a quarter of the emission rate from sources in North America and Europe. In the case of NH_x, 258 Moles-N m^–2 yr^–1 (3.6 Tg N) are deposited to the NAO and the Caribbean; this is about 6% of the global continental emissions. There is relatively little data on the deposition rate of organic nitrogen species; nonetheless, this evidence suggests that concentrations and deposition rates are comparable to those for inorganic nitrogen.Because of anthropogenic emissions, the present-day deposition rate of NO_y to the NAO is about five times greater than pre-industrial times largely due to emissions from energy production and biomass burning. The present-day emissions of NH_x from continental anthropogenic sources are about four-to-five times greater than natural sources, mostly due to the impact of emissions from animal wastes associated with food production. Indeed, present-day emissions of NH_x from animal waste are estimated to be about 10 times greater than the pre-human era. The deposition rate of mineral dust to the NAO is about 170 Tg yr^–1; deposited with the dust (assuming average crustal abundances) is about 6 Tg yr^–1 of Fe and 0.2 Tg yr^–1 of P. Dust deposition in the NAO is almost completely attributable to transport from North African sources; a substantial fraction of the dust over the NAO is probably mobilized as a consequence of land use practices in arid regions and, consequently, it should be regarded as a pollutant.     

Nitrogen and phosphorus budgets of the North Atlantic Ocean and its watershed

Anthropogenic food and energy production extensively mobilize reactive nitrogen (N) in the watershed of the North Atlantic Ocean (NAO). There is wide spread N distribution by both hydrologic and atmospheric processes within the watershed of the NAO, resulting in reactive N accumulation in terrestrial systems. Net denitrification in most estuaries and continental shelves exceeds the amount of N supplied to the shelves by rivers and requires a supply of nitrate from the open ocean. Thus riverine N is only transported to the open ocean in a few areas with the flow from a few major rivers (e.g., Amazon). Atmospheric N deposition to the open ocean has increased and may increase the productivity of the surface ocean. In addition, as a consequence of increased Fe deposition to the open ocean (due in part to anthropogenic processes), the rate of biological N-fixation may have increased resulting in N accumulation in the ocean. Phosphorus (P) is also mobilized by anthropogenic processes (primarily food production). Relative to N, more of the P is transported across the shelf to the open ocean from both estuaries and major rivers. There are several consequences of the increased availability of N and P that are unique to each element. However, the control on primary productivity in both coastal and open ocean ecosystems is dependent on a complex and poorly understood interaction between N and P mobilization and availability.     

Nitrogen fixation in subarctic streams influenced by beaver (Castor canadensis)

Nitrogen fixation was measured in four subarctic streams substantially modified by beaver (Castor canadensis) in Quebec. Acetylene-ethylene (C₂H₂ C₂H₄) reduction techniques were used during the 1982 ice-free period (May–October) to estimate nitrogen fixation by microorganisms colonizing wood and sediment. Mean seasonal fixation rates were low and patchy, ranging from zero to 2.3 × 10^–3 µmol C₂H₄ · cm^–2 · h^–1 for wood, and from zero to 7.0 × 10^–3 µmol C₂H₄ · g AFDM^–1 · h^–1 for sediment; 77% of all wood and 63% of all sediment measurements showed no C₂H₂ reduction. Nonparametric statistical tests were unable to show a significant difference (p > 0.05) in C₂H₂ reduction rates between or within sites for wood species or by sediment depth.Nitrogen contributed by microorganisms colonizing wood in riffles of beaver influenced watersheds was small (e.g., 0.207 g N · m^–2 · y^–1) but greater than that for wood in beaver ponds (e.g., 0.008 g N · m^–2 · y^–1) or for streams without beaver (e.g., 0.003 g N · m^–2 · y^–1). Although mass specific nitrogen fixation rates did not change significantly as beaver transform riffles into ponds, the nitrogen fixed by organisms colonizing sediment in pond areas (e.g., 5.1 g N · m^–2 · y^–1) was greater than that in riffles (e.g., 0.42 g N · m^–2 · y^–1). The annual nitrogen contribution is proportional to the amount of sediment available for microbial colonization. We estimate that total nitrogen accumulation in sediment, per unit area, is enhanced 9 to 44 fold by beaver damming a section of stream.     

Turbidity enhances feeding abilities of larval Pacific herring,Clupea harengus pallasi

Fishes inhabiting estuaries, rivers, and embayments are subject to turbid conditions. Larvae of many fishes utilize estuaries as nursery areas. For visual plankton feeders such as larval fishes, turbidity may reduce search and reaction distances, resulting in lowered feeding abilities. In this study feeding Pacific herring larvae, Clupea harengus pallasi, were exposed to suspensions of estuarine sediment and Mount Saint Helens volcanic ash at concentrations ranging from 0 mg · l^–1 to 8 000 mg · l^–1. In all experiments, maximum feeding incidence and intensity occurred at levels of suspension of either 500 mg · l^–1 or 1000 mg · l^–1 with values significantly greater than controls (0 mg · l^–1). Feeding decreased at greater concentrations. The suspensions may enhance feeding by providing visual contrast of prey items on the small perceptive scale used by the larvae. Larval residence in turbid environments such as estuaries may serve to reduce predation from larger, visual planktivores, while searching ability in the small larval perceptive field is not decreased.     

Evaluation of the nutrient inputs to a coastal lagoon: the case of the Ria de Aveiro,Portugal

The Ria de Aveiro estuary-coastal lagoon system of northern Portugal is estimated to currently receive mean annual influxes of total nitrogen (N) and total phosphorus (P) of c. 6118 t y^–1 and 779 t y^–1, respectively, from its influent rivers. In low summer flows the mean N and P fluxes decrease to c. 10% of the annual average. The sewage contribution to the inland-derived N load on an annual basis is c. 5% but, during the summer low flow conditions, the sewage component increases to c. 65% of the total river loading. The sewage contribution to the inland-derived P load on an annual basis is c. 11% but, during the dry season, it is 1.2 times larger than the river-derived flux. The construction of a regional sewer system linked to a submarine outfall, due for completion in 2005, is expected to lead to a reduction in nutrient fluxes from inland to the lagoon of c. 15% for N and c. 26% for P relative to the present values. While this system will reduce the nutrient loading in the upper reaches of the lagoon, an increase in nutrients derived from the ocean is anticipated, due to the proximity of the outfall to the inlet.     

Postsynaptic effects of substance P in frog neuromuscular junction

We have studied the effect of substance P on the end-plate currents (EPC) and the miniature EPC (MEPC) after acetylcholine esterase (ACE) inhibition in the cut neuromuscular preparation of the frog sartorius muscle using the voltage-clamp technique. At concentrations of 5·10^–7–1·10^–6 moles/liter substance P had no effect on the amplitude and the time characteristics of single EPC and MEPC but promoted prolongation of EPC decay on repetitive stimulation of the nerve with a frequency of 10/sec, indicating intensification of postsynaptic potentiation. Elevation of the concentration of the given peptide to 5·10^–6 moles/liter led to the shortening of the decay of single EPC and a more marked depression of the EPC amplitude in the trains as compared to the control, reflecting a decrease in the sensitivity of the postsynaptic membrane to the mediator, i.e., development of desensitization.S. V. Kurashov State Medical Institute, Kazan. I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Leningrad. Translated from Neirofiziologiya, Vol. 23, No. 4, pp. 436–441, July–August, 1991.     

Bacterial productivity and microbial biomass in tropical mangrove sediments

Bacterial productivity (³H-thymidine incorporation into DNA) and intertidal microbenthic communities were examined within five mangrove estuaries along the tropical northeastern coast of Australia. Bacteria in mangrove surface sediments (0–2 cm depth) were enumerated by epifluorescence microscopy and were more abundant (mean and range: 1.1(0.02–3.6)×10¹¹ cells·g DW^–1) and productive (mean: 1.6 gC·m^–2· d^–1) compared to bacterial populations in most other benthic environments. Specific growth rates (¯x=1.1) ranged from 0.2–5.5 d^–1, with highest rates of growth in austral spring and summer. Highest bacterial numbers occurred in winter (June–August) in estuaries along the Cape York peninsula north of Hinchinbrook Island and were significantly different among intertidal zones and estuaries. Protozoa (10⁵–10⁶·m^–2, pheopigments (0.0–24.1g·gDW^–1) and bacterial productivity (0.2–5.1 gC·m^–2·d^–1) exhibited significant seasonality with maximum densities and production in austral spring and summer. Algal biomass (chlorophylla) was low (mean: 1.6g·gDW^–1) compared to other intertidal sediments because of low light intensity under the dense forest canopy, especially in the mid-intertidal zone. Partial correlation analysis and a study of possible tidal effects suggest that microbial biomass and bacterial growth in tropical intertidal sediments are regulated primarily by physicochemical factors and by tidal flushing and exposure. High microbial biomass and very high rates of bacterial productivity coupled with low densities of meiofaunal and macroinfaunal consumers observed in earlier studies suggest that microbes may be a sink for carbon in intertidal sediments of tropical mangrove estuaries.     

The role of rock weathering in the phosphorus budget of terrestrial watersheds

Residual soils (saprolites) developed on crystalline rocks appear to form by an essentially isovolumetric process (i.e. without dilation or compaction). Isovolumetric geochemical analysis of a suite of saprolite samples developed on a common parent rock can be used to estimate the relative rates of long-term losses of P and Si during weathering. Using the export of dissolved Si in rivers as a weathering index, one can then estimate the rate of P release due to chemical weathering by means of the P-Si loss ratio in saprolite. For three basins where data are available (Liberty Hill, SC; Amazon River, Brazil: Rio Negro, Brazil) estimated P weathering release rates are 163, 457, and 242 moles P km^–2 yr^–1 respectively. These compare to precipitation inputs of 684, 700 and 630 moles P km^–2 yr^–1 and total river exports of 256, 4490 and 820 moles P km^–2 yr^–1, respectively. The Rio Negro shows a near perfect balance between the input of P via precipitation and chemical weathering and the riverine output of dissolved and suspended P. This system, however, raised the unsolved problem of the source that supports the atmospheric P input.     

The role of cattle in the volatile loss of nitrogen from a shortgrass steppe

The cycling and volatile loss of N derived from cattle urine at upland and lowland sites within the shortgrass steppe of eastern Colorado was studied, using¹⁵N-labelled urea as an N source. Losses of NH ₀ ³ were determined by direct measurement and by difference. Losses were higher from coarse (27% summer, 12% winter) than from fine textured (0–2%) soils. Immobilization and plant uptake of N accounted for significant amounts of added N. Extrapolating our plot measurements to a typical pasture, using spatially and temporally stratified urine deposition data, losses from upland sites were calculated to be 0.016 g N · m^-2 · y^-1, while losses from lowland sites were negligible. This resulted in an average loss of 0.011 g N · m^-2 · y^-1 for a pasture divided 70:30 between uplands and lowlands. The loss of urine N calculated assuming no spatial stratification would be sevenfold higher (0.076 g N · m^-2 · y^-1). Losses of NH ₀ ³ from urine, animal biomass removal, and NH₂O loss totaled only 0.07 g N · m^-2 · y^-1 , or about 25% of wet deposition input. We calculated a potential loss of NH ₀ ³ from senescing vegetation of 0.26 g N · m^-2 · y^-1, an order of magnitude larger than all other losses combined.     

Bacterial productivity in the water column and sediments of the Georgia (USA) coastal zone: Estimates via direct counting and parallel measurement of thymidine incorporation

Three methods of estimating bacterial productivity were compared using parallel samples of Atlantic Ocean water (within 0.25–15 km of the Georgia coast). The frequency-of-dividing cells (FDC) method and the [³H]thymidine incorporation method gave results which were strongly correlated (r=0.97), but the FDC estimates were always higher (X2 to X7) than the [³H]thymidine estimates. Estimates of bacterial productivity ranged from 2–4×10⁸ cells·l^–1·h^–1 at 0.25 km from shore to 1–9×10⁷cells·l^–1·h^–1 at 15 km. A method involving incubation of 3-m filtrates and direct counting gave results that could not be easily translated into estimates of bacterial productivity. Application of the FDC method to sediment samples gave high productivity estimates, which could be not reconciled with productivity estimates based on sediment oxygen uptake.     

The North Atlantic subpolar gyre and the marine migration of Atlantic salmon Salmo salar: the ‘Merry‐Go‐Round’ hypothesis

One model for marine migration of Atlantic salmon Salmo salar proposes that North American and southern European stocks (<62° N) move directly to feeding grounds off west Greenland, then overwinter in the Labrador Sea, whereas northern European stocks (>62° N) utilize the Norwegian Sea. An alternate model proposes that both North American and European stocks migrate in the North Atlantic Subpolar Gyre (NASpG) where S. salar enter the NASpG on their respective sides of the Atlantic, and travel counterclockwise within the NASpG until returning to natal rivers. A review of data accumulated during the last 50 years suggests a gyre model is most probable. Freshwater parr metamorphose into smolts which have morphological, physiological and behavioural adaptations of epipelagic, marine fishes. Former high‐seas fisheries were seasonally sequential and moved in the direction of NASpG currents, and catches were highest along the main axis of the NASpG. Marking and discrimination studies indicate mixed continental origin feeding aggregations on both sides of the Atlantic. Marked North American smolts were captured off Norway, the Faroe Islands, east and west Greenland, and adults tagged at the Faroes were recovered in Canadian rivers. Marked European smolts were recovered off Newfoundland and Labrador, west and east Greenland, and adults tagged in the Labrador Sea were captured in European rivers. High Caesium‐137 (¹³⁷Cs) levels in S. salar returning to a Quebec river suggested 62·3% had fed at or east of Iceland, whereas levels in 1 sea‐winter (SW) Atlantic Canada returnees indicated 24·7% had fed east of the Faroes. Lower levels of ¹³⁷ Cs in returning 1SW Irish fish suggest much of their growth occurred in the western Atlantic. These data suggest marine migration of S. salar follows a gyre model and is similar to other open‐ocean migrations of epipelagic fishes.     

Sedimentation rates in a Swiss-Italian lake measured with sediment traps

Tripton sedimentation was investigated in the eutrophic Lake Lugano (Ponte Tresa basin) from October 1979 to October 1980. The annual amount of tripton collected was 748 g · m^–2 · y^–1. Phosphorus, nitrogen and organic carbon fluxes into the hypolimnion were estimated to be 1.9, 16.2 and 121 g · m^–2y^–1 respectively. Mineralization rates into the trophogenic layer varied from 11% to 19% per day during summer stratification. The regeneration processes contribute about 60% of the calculated P deficit in the epilimnion. The tripton is decomposed mostly in the metalimnion, out of the euphotic zone; from here the phosphorus is carried back to the overlying waters by diffusion processes.     

Chaff piles of harvester ant (Messor spp.) nests in a desert ecosystem

Summary The plant species composition of the chaff piles of three species of harvester ant (Messor spp.) and the contribution of the chaff to the organic pool were studied from August 1985 to July 1987. There were distinct differences in the plant species composition of the chaff of the three species. We attribute this to the different diets of the three species, which reflect the relative sizes of their individuals and their foraging strategies. The amount of chaff accumulated varies greatly between the three species (Messor rugossus: 127–196 g · ha^–1 · y^–1;Messor ebeninus: 2823–4437 g · ha^–1 · y^–1;Messor arenarius: 2165–2535 g · ha^–1 · y^–1), although the number of nests per hectare is virtually the same. We found that the amount of chaff is related to the rate of activity and the size of the individuals of each of the three ant species. The total chaff accumulated during the study period was 19.2 kg · ha^–1, which is an important contribution to the organic matter in the soil in the Negev desert ecosystem.     

Spatial and temporal variability of dissolved sulfur compounds in European estuaries

The spatial and temporal distribution of Dimethylsulfide (DMS),Dimethylsulfoniopropionate (DMSP) – its precursor –,Dimethylsulfoxyde (DMSO) – one of its oxidation products –Carbonyl sulfide (COS) and Carbonyl disulfide (CS₂) wereinvestigated during nine oceanographic cruises carried out in six majortidal European estuaries between July 1996 and May 1998. Low levels ofDMS were recorded (mean of 0.6 nM, standard deviation () 0.3 nM)during these 9 cruises. Positive correlations between DMS and salinitywere frequently observed, with the highest DMS concentrations in theplume of estuaries, which could be explained by change in phytoplanktonspeciation from estuarine to shelf waters. Strong correlation betweenDMSP and DMS was reported for most of the estuaries denoting a simpleconservative mixing of riverine and marine waters controlled by tide. Incontrast to DMS, significant levels of COS and CS₂ with meanconcentrations of 220 ± 150 (pM; pM =10^–12 M) and 25 ± 6 pM respectively wererecorded in four estuaries, indicating that estuaries could be asignificant source of these compounds.     

Phosphate in the sediments of the Gulf of Lions (NW Mediterranean Sea), relationship with input by the river Rhone

We investigated P-input by the Rhone river into the Mediterranean Sea taking into account P trapped in the surface sediment of the Gulf of Lions. Total phosphate concentration was determined every cm in the upper 10 cm-layer of sediments sampled at 11 stations in the Gulf of Lions during two cruises (March 1998 and January 1999). Two low downward gradients, one East–West and another North–South, with distance to the Rhone river mouth were found. Except at one station, total phosphate concentration in surface sediments was found to be constant with depth down to 10 cm. Values for individual stations ranged between 400 and 700 g g^–1 with an average value of 547 g g^–1 (st. dev. = 63 g g^–1) for the whole gulf. The low variability in total-P concentration in sediments is in contrast to the large variability in suspended matter load of the river Rhone and suggests the dominance of authigenic P removal mechanisms in P burial. The total P-pool in the upper 10 cm-layer of the sediments in the gulf was estimated at 562 kt, with about 80% trapped into the shelf and 20% into the slope. Annual P-deposition was estimated as 7.2–12.4 kt y^–1, from the P-pool in the sediment and the sedimentation rates. This is equivalent to a previous estimation of the river Rhone input, estimated to be about 6.5–12.2 kt y^–1. As the Rhone is the major river flowing into the Mediterranean Sea, total P in surface sediments of the Gulf of Lions should be taken into account in P-budgets at the scale of the Mediterranean Sea.     

Areas of shallow water in the North Atlantic

We report here estimates of the areas of water that are between 1 and 100 m and between 1 and 200 m deep in the North Atlantic, including the Caribbean Sea, the Gulf of Mexico, Hudson Bay and the North Sea, but excluding estuaries. The total areas within these depths, from the equator to 70° N, are 3.91 × 10⁶ km² and 5.66 × 10⁶ km², respectively. We also report the respective areas by selected geographic regions.     

Longitudinal Spatial Patterns of Bacterial Production and Respiration in a Large River–Estuary: Implications for Ecosystem Carbon Consumption

Rivers and estuaries transport organic carbon (C) from terrestrial and freshwater ecosystems to the marine environment. During this transit, bacteria actively utilize and transform organic C, but few studies have measured detailed spatial variation in rates of bacterial respiration (BR) and production (BP). We measured BP at 39 stations and BR at 12 stations at monthly intervals along a 200-km reach of the tidal Hudson River. We observed strong repeatable spatial patterns for both BP and BR, with rates declining in the downstream direction. Bacterial Production had much greater dynamic range of spatial variation than BR. We used the detailed seasonal and spatial data on BP and BR to measure the total C demand of bacteria at several scales. We calculated volumetric and areal rates for 12 sections of the Hudson, as well as the total C utilization. Volumetric BR averaged 20 g-C-m^–3 y^–1, but it was highest in the most upstream section at 30 g C m^–3 y^–1. Areal rates averaged over the entire river were 174 g C m^–2 y^–1, but they were 318 g C m^–2 y^–1 in the deepest section of the river, indicating the importance of morphometric variation. Total bacterial C demand increased downriver with increasing total volume. Overall, bacteria in the freshwater section of the river consumed approximately 18–25.5 × 10⁹ g C y^–1, about 20% of the total organic C load.     

The nitrogen cycle in lodgepole pine forests,southeastern Wyoming

Storage and flux of nitrogen were studied in several contrasting lodgepole pine (Pinus contorta spp.latifolia) forests in southeastern Wyoming. The mineral soil contained most of the N in these ecosystems (range of 315–860 g · m^–2), with aboveground detritus (37.5–48.8g · m^–2) and living biomass (19.5–24.0 g · m^–2) storing much smaller amounts. About 60–70% of the total N in vegetation was aboveground, and N concentrations in plant tissues were unusually low (foliage = 0.7% N), as were N input via wet precipitation (0.25 g · m^–2 · yr^–1), and biological fixation of atmospheric N (<0.03 g · m^–2 · yr^–1, except locally in some stands at low elevations where symbiotic fixation by the leguminous herbLupinus argenteus probably exceeded 0.1 g · m^–2 · yr^–1).Because of low concentrations in litterfall and limited opportunity for leaching, N accumulated in decaying leaves for 6–7 yr following leaf fall. This process represented an annual flux of about 0.5g · m^–2 to the 01 horizon. Only 20% of this flux was provided by throughfall, with the remaining 0.4g · m^–2 · yr^–1 apparently added from layers below. Low mineralization and small amounts of N uptake from the 02 are likely because of minimal rooting in the forest floor (as defined herein) and negligible mineral N (< 0.05 mg · L^–1) in 02 leachate. A critical transport process was solubilization of organic N, mostly fulvic acids. Most of the organic N from the forest floor was retained within the major tree rooting zone (0–40 cm), and mineralization of soil organic N provided NH₄ for tree uptake. Nitrate was at trace levels in soil solutions, and a long lag in nitrification was always observed under disturbed conditions. Total root nitrogen uptake was calculated to be 1.25 gN · m^–2 · yr^–1 with estimated root turnover of 0.37-gN · m^–2 · yr^–1, and the soil horizons appeared to be nearly in balance with respect to N. The high demand for mineralized N and the precipitation of fulvic acid in the mineral soil resulted in minimal deep leaching in most stands (< 0.02 g · m^–2 · yr^–1). These forests provide an extreme example of nitrogen behavior in dry, infertile forests.