The biogeochemistry of nitrogen in freshwater wetlands

The biogeochemistry of N in freshwater wetlands is complicated by vegetation characteristics that range from annual herbs to perennial woodlands; by hydrologic characteristics that range from closed, precipitation-driven to tidal, riverine wetlands; and by the diversity of the nitrogen cycle itself. It is clear that sediments are the single largest pool of nitrogen in wetland ecosystems (100's to 1000's g N m^-2) followed in rough order-of-magnitude decreases by plants and available inorganic nitrogen. Precipitation inputs (< 1–2 g N m^-2 yr^-1) are well known but other atmospheric inputs, e.g. dry deposition, are essentially unknown and could be as large or larger than wet deposition. Nitrogen fixation (acetylene reduction) is an important supplementary input in some wetlands (< < 1–3 g N m^-2 yr^-1) but is probably limited by the excess of fixed nitrogen usually present in wetland sediments.Plant uptake normally ranges from a few g N m^-2 yr^-1 to 35 g N m^-2 yr^-1 with extreme values of up to 100g N m^-2 yr^-1 Results of translocation experiments done to date may be misleading and may call for a reassessment of the magnitude of both plant uptake and leaching rates. Interactions between plant litter and decomposer microorganisms tend, over the short-term, to conserve nitrogen within the system in immobile forms. Later, decomposers release this nitrogen in forms and at rates that plants can efficiently reassimilate.The NO₃ formed by nitrification (< 0.1 to 10 g N m^-2 yr^-1 has several fates which may tend to either conserve nitrogen (uptake and dissimilatory reduction to ammonium) or lead to its loss (denitrification). Both nitrification and denitrification operate at rates far below their potential and under proper conditions (e.g. draining or fluctuating water levels) may accelerate. However, virtually all estimates of denitrification rates in freshwater wetlands are based on measurements of potential denitrification, not actual denitrification and, as a consequence, the importance of denitrification in these ecosystems may have been greatly over estimated.In general, larger amounts of nitrogen cycle within freshwater wetlands than flow in or out. Except for closed, ombrotrophic systems this might seem an unusual characteristic for ecosystems that are dominated by the flux of water, however, two factors limit the opportunity for N loss. At any given time the fraction of nitrogen in wetlands that could be lost by hydrologic export is probably a small fraction of the potentially mineralizable nitrogen and is certainly a negligible fraction of the total nitrogen in the system. Second, in some cases freshwater wetlands may be hydrologically isolated so that the bulk of upland water flow may pass under (in the case of floating mats) or by (in the case of riparian systems) the biotically active components of the wetland. This may explain the rather limited range of N loading rates real wetlands can accept in comparison to, for example, percolation columns or engineered marshes.     

Benithic-pelagic coupling of nutrient metabolism along an estuarine eutrophication gradient

The shallow, brackish (11–18% salinity) Roskilde Fjord represents a eutrophication gradient with annual averages of chlorophyll, ranging from 3 to 25 mg chl a m^–3. Nutrient loadings in 1985 were 11.3–62.4 g N m^–2 yr^–1 and 0.4–7.3 g P m^–2 yr^–1. A simple one-layer advection-diffusion model was used to calculate mass balances for 7 boxes in the fjord. Net loss rates varied from –32.2 to 17.9 g P m^–2 yr^–1 and from –3.3 to 66.8 g N m^–2, corresponding to 74% of the external P-loading and 88% of the external N-loading to the entire estuary.Gross sedimentation rates measured by sediment traps were between 7 and 52 g p m^–2 yr^–1 and 50 and 426 g N M^–2 yr^–1, respectively. Exchangeable sediment phosphorus varied in annual average between 2.0 and 4.8 g P m^–2 and exchangeable sediment nitrogen varied from 1.9 to 33.1 g N m–1. Amplitudes in the exchangeable pools followed sedimentation peaks with delays corresponding to settling rates of 0.3 m d^–1. Short term nutrient exchange experiments performed in the laboratory with simultaneous measurements of sediment oxygen uptake showed a release pattern following the oxygen uptake, the changes in the exchangeable pools and the sedimentation peaks.The close benthic-pelagic coupling also exists for the denitrification with maxima during spring of 5 to 20 mmol N m^–2 d–1. Denitrification during the nitrogen-limited summer period suggests dependence on nitrification. Comparisons with denitrification from other shallow estuaries indicate a maximum for denitrification in estuaries of about 250 µmol N m^–2 h^–2 achieved at loading rates of about 25–125 g N m^–2 yr^–1.     

The impacts of re-introducing Mississippi River water on the hydrologic budget and nutrient inputs of a deltaic estuary

Most wetlands of the Mississippi deltaic plain are isolated from riverine input due to flood control levees along the Mississippi River. These levees have altered hydrology and ecology and are a primary cause of massive wetland loss in the delta. River water is being re-introduced into coastal basins as part of a large-scale ecological engineering effort to restore the delta. We quantified freshwater, nitrogen, and phosphorus inputs to the Breton Sound Estuary for three climatically different years (2000, 2001, and 2002). Water budgets included precipitation, potential evapotranspiration, the diversion, stormwater pumps, and groundwater. Precipitation contributed 48–57% of freshwater input, while the diversion accounted for 33–48%. Net groundwater input accounted for less than 0.05% of freshwater inputs. Inputs of ammonium (NH₄-N), nitrate (NO₃-N), total nitrogen (TN), and total phosphorus (TP) were determined for each of the water sources. Atmospheric deposition was the most important input of NH₄-N (57–62% or 1.44 × 10⁵–2.32 × 10⁵ kg yr⁻¹) followed by the diversion. The diversion was the greatest source of NO₃-N (67–83%, 7.78 × 10⁵–1.64 × 10⁶ kg yr⁻¹) and TN (60–71%). The diversion contributed 41–60% of TP input (1.17 × 10⁵–2.32 × 10⁵ kg yr⁻¹). Annual loading rates of NH₄-N and NO₃-N were 0.17–0.27 and 1.2–2.3 g N m⁻² yr⁻¹, respectively, for the total basin indicating strong retention of nitrogen in the basin. Nitrogen retention through denitrification and burial was estimated for the upper basin.     

Predictive models for phosphorus retention in wetlands

The potential of wetlands to efficiently remove (i.e., act as a nutrient sink) or to transform nutrients like phosphorus under high nutrient loading has resulted in their consideration as a cost-effective means of treating wastewater on the landscape. Few predictive models exist which can accurately assess P retention capacity. An analysis of the north American data base (NADB) allowed us to develop a mass loading model that can be used to predict P storage and effluent concentrations from wetlands. Phosphorus storage in wetlands is proportional to P loadings but the output total phosphorus (TP) concentrations increase exponentially after a P loading threshold is reached. The threshold P assimilative capacity based on the NADB and a test site in the Everglades is approximately 1 g m^–2 yr^–1. We hypothesize that once loadings exceed 1 g m^–2 yr^–1 and short-term mechanisms are saturated, that the mechanisms controlling the uptake and storage of P in wetlands are exceeded and effluent concentrations of TP rise exponentially. We propose a One Gram Rule for freshwater wetlands and contend that this loading is near the assimilative capacity of wetlands. Our analysis further suggests that P loadings must be reduced to 1 g m^–2 yr^–1 or lower within the wetland if maintaining long-term low P output concentrations from the wetlands is the central goal. A carbon based phosphorus retention model developed for peatlands and tested in the Everglades of Florida provided further evidence of the proposed One Gram Rule for wetlands. This model is based on data from the Everglades areas impacted by agricultural runoff during the past 30 years. Preliminary estimates indicate that these wetlands store P primarily as humic organic-P, insoluble P, and Ca bound P at 0.44 g m^–2 yr^–1 on average. Areas loaded with 4.0 g m^–2 yr^–1 (at water concentrations>150 g·L^–1 TP) stored 0.8 to 0.6 g m^–2 yr^–1 P, areas loaded with 3.3 g m^–2 yr^–1 P retained 0.6 to 0.4 g m^–2 yr^–1 P, and areas receiving 0.6 g m^–2 yr^–1 P retained 0.3 to 0.2 g m^–2 yr^–1. The TP water concentrations in the wetland did not drop below 50 g·L^–1 until loadings were below 1 g m² yr^–1 P.     

Retention of nutrients in river basins

In Denmark, as in many other European countries, the diffuse losses of nitrogen (N) and phosphorus (P) from the rural landscape are the major causes of surface water eutrophication and groundwater pollution. The export of total N and total P from the Gjern river basin amounted to 18.2 kg ha^–1 and 0.63 kg P ha^–1 during June 1994 to May 1995. Diffuse losses of N and P from agricultural areas were the main nutrient source in the river basin contributing 76% and 51%, respectively, of the total export.Investigations of nutrient cycling in the Gjern river basin have revealed the importance of permanent nutrient sinks (denitrification and overbank sedimentation) and temporary nutrient storage in watercourses. Temporary retention of N and P in the watercourses thus amounted to 7.2–16.1 g N m^–2 yr^–1 and 3.7–8.3 g P m^–2 yr–1 during low-flow periods. Deposition of P on temporarily flooded riparian areas amounted from 0.16 to 6.50 g P m^–2 during single irrigation and overbank flood events, whereas denitrification of nitrate amounted on average to 7.96 kg N yr^–1 per running metre watercourse in a minerotrophic fen and 1.53 kg N yr^–1 per linear metre watercourse in a wet meadow. On average, annual retention of N and P in 18 Danish shallow lakes amounted to 32.5 g N m^–2 yr^–1 and 0.30 g P m^–2 yr^–1, respectively, during the period 1989–1995.The results indicate that permanent nutrient sinks and temporary nutrient storage in river systems represent an important component of river basin nutrient budgets. Model estimates of the natural retention potential of the Gjern river basin revealed an increase from 38.8 to 81.4 tonnes yr^–1 and that P-retention increased from –0.80 to 0.90 tonnes yr^–1 following restoration of the water courses, riparian areas and a shallow lake. Catchment management measures such as nature restoration at the river basin scale can thus help to combat diffuse nutrient pollution.     

A first generation ecosystem model of the Des Plaines River experimental wetlands

A first generation wetland ecosystem model is developed from first-year field data for four constructed freshwater marshes at the Des Plaines River Wetland Demonstration Project in northeastern Illiois, USA. The model, which includes hydrology, sediment and phophorus algorithms common for all four wetlands, was used for the integration of data collected as part of the many different aspects of the demonstration project and for prediction of wetland function under varying hydrologic and chemical loadings. Stepwise calibration was completed on the model for the hydrology, sediment, productivity, and phosphorus submodels. Separate sedimentation coefficients were necessary for each wetland land and each season. Preliminary simulations investigate the role of changing flow conditions in the wetlands on phosphorus retention. As simulated inflow increased from 34 to 136 cm/wk, phosphorus retention increased from 1.5 to 3.3 g P m⁻² yr⁻¹ while retention efficiency decreased from 67 to 37%.     

Carbon and nitrogen cycling in intertidal sediments near Doel,Scheldt Estuary

Carbon and nitrogen cycling in intertidal mud flat sediments in the Scheldt Estuary was studied using measurements of carbon dioxide, methane and nitrous oxide emission rates and pore-water profiles of CO₂, ammonium and nitrate. A comparison between chamber measured carbon dioxide fluxes and those based on CO₂ pore-water gradients using Fick's First law indicates that apparent diffusion coefficients are 2 to 28 times higher than bulk sediment diffusion coefficients based on molecular diffusion. Seasonal changes in gaseous carbon fluxes or CO₂ pore water concentrations cannot be used directly, or in a simple way, to determine seasonal rates of mineralization, because of marked seasonal changes in pore-water storage and exchange parameters.The annual amount of carbon delivered to the sediment is 42 mol m^–2, of which about 42% becomes buried, the remaining being emitted as methane (7%) or carbon dioxide (50%). Each year about 2.6 mol N m^–2 of particulate nitrogen reaches the sediment; 1.1 mol m^–2 is buried and 1.6 mol m^–2 is mineralized to ammonium. Only 0.42 mol m^–2 yr^–1 of the ammonium produced escapes from the sediments, the remaining being first nitrified (1.2 mol m^–2 yr^–1) and then denitrified (1.7 mol m^–2 yr^–1). Simple calculations indicate that intertidal sediments may account for about 14% and 30% of the total estuarine retention of nitrogen and carbon, respectively.     

Release of sedimentary nitrogen and phosphorus in polder ditches of a low-moor peat area

The release of N and P from the sediment of two ditches, one (A) dominated by filamentous algae and the other (B) by water-lilies, was estimated by core and enclosure experiments. The release rates for ditch A tended to be higher than those for ditch B. Sediment cores covered by a filamentous algae layer released about 1.5 times more N and P than those from which the layer had been removed. During the incubation of the cores in the dark at 20°C for 2–3 weeks, about 10% of the N in the filamentous algae layer was mineralized. The mineralization could be described as a first-order reaction with a rate constant of about 0.2 d^–1. On average the cores of ditches A and B released about 40 mg mineral N and 3 mg.m^–2.d^–1 soluble reactive phosphorus. Defining the release from the sediment in the enclosures as the net increase of N and P in the water phase and in the vegetation minus the input, a negative net release,i.e. net accumulation of N and P in the sediment, was found over the summer half of the year. The negative values were due to the significant N and P input, resulting from pumping ditch water into the enclosures in order to compensate for downward seepage. From the enclosure experiments a downward seepage rate of 14 mm.d^–1 and an external load of about 6 g.m^–2 total N and 0.6 g.m^–2 total P during the summer half of the year —i.e. 33 mg.m^–2.d^–1 N and 3 mg.m^–2.d^–1 P. respectively — was calculated for the ditches. Tentative gross release rates — based on the sum of the positive net release of N and P into the water phase over 1–2 weeks intervals and the net increase of N and P in the vegetation — converted to 20°C and allowing for underestimation of the primary production by a factor of 5, amounted to 58 mg mineral N and 7 mg.m^–2.d^–1 soluble reactive phosphorus during the summer half of the year. Combining the rates estimated by cores and enclosures and converting them to rates at the mean water temperature during the summer half of the year, the release of mineral N and soluble reactive phosphorus roughly amounted to 40 and 4 mg.m^–2.d^–1, respectively. The release rates as well as the external load indicated a relatively low eutrophication of the ditches.     

Phosphorus and nitrogen retention in five Precambrian shield wetlands

Phosphorus and nitrogen mass balances of five wetlands (two beaver ponds, two conifer-Sphagnum swamps and one sedge fen) situated in three catchments in central Ontario, Canada, were measured. Monthly and annual input-output budgets of total phosphorus (TP), total nitrogen (TN), total organic nitrogen (TON), total inorganic nitrogen (TIN), ammonium ion (NH₄ ⁺ -N), nitrate (NO ₃ ^– -N) and dissolved organic carbon (DOC) were estimated for the five wetlands during the 1982–83 and 1983–84 water years. Except for the deepest beaver pond (3.2 m) which had annual TP retention of –44% (–0.030 ± 0.015 g m^–2 yr^–1), the wetlands retained < 0.001 to 0.015 g M^–2 yr^–1 ; however, this wasless than 20% of the inputs and the estimated budget uncertainties were equal to or greater than the retention rates. Annual TN retentions ranged from –0.44 to 0.56 g m^–2 yr^–1 (–12 to 4%) but were not significantly different from zero. The wetlands transformed nitrogen by retaining TIN (16 to 80% RT) and exporting an equivalent amount as TON (–7 to 102% RT). The beaver ponds, however, retained NO ₃ ^– while NH ₄ ⁺ was passed through or the outputs exceeded the inputs. In contrast, the conifer swamps retained both NH ₄ ⁺ and NO ₃ ^– . DOC fluxes into and out of the beaver ponds were equal (–18 and 4% RT) but output from the conifer swamps exceeded input by > 90%. Marked seasonal trends in nutrient retention were observed. Nutrient retention coincided with low stream flow, increased evapotranspiration and biotic uptake during the summer. Net nutrient export occurred during the winter and spring when stream flows were highest and biotic uptake was low.     

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.     

Interactions between sediment and water in a shallow and hypertrophic lake: a study on phytoplankton collapses in Lake Søbygård,Denmark

Short-term changes in phytoplankton and zooplankton biomass have occurred 1–3 times every summer for the past 5 years in the shallow and hypertrophic Lake Søbygård, Denmark. These changes markedly affected lake water characteristics as well as the sediment/water interaction. Thus during a collapse of the phytoplankton biomass in 1985, lasting for about 2 weeks, the lake water became almost anoxic, followed by rapid increase in nitrogen and phosphorus at rates of 100–400 mg N M^–2 day^–1 and 100–200 mg P m^–1 day^–1. Average external loading during this period was about 350 mg N m^–2 day^–1 and 5 mg P m^–2 day^–1, respectively.Due to high phytoplankton biomass and subsequently a high sedimentation and recycling of nutrients, gross release rates of phosphorus and nitrogen were several times higher than net release rates. The net summer sediment release of phosphorus was usually about 40 mg P m^–2 day^–1, corresponding to a 2–3 fold increase in the net phosphorus release during the collapse. The nitrogen and phosphorus increase during the collapse is considered to be due primarily to a decreased sedimentation because of low algal biomass. The nutrient interactions between sediment and lake water during phytoplankton collapse, therefore, were changed from being dominated by both a large input and a large sedimentation of nutrients to a dominance of only a large input. Nitrogen was derived from both the inlet and sediment, whereas phosphorus was preferentially derived from the sediment. Different temperature levels may be a main reason for the different release rates from year to year.     

Primary production, nutrient dynamics, and accretion of a coastal freshwater forested wetland assimilation system in Louisiana

This study reports on the response of a tidal, freshwater forested wetland ecosystem to long-term input of secondarily treated municipal effluent from the City of Mandeville, LA. Measurements of hydrology, nutrients, and aboveground net primary productivity were made from September 1998 through March 2002. Accretion measurements were made in October 2000 and October 2004. The major hydrologic inputs to the system were the effluent, precipitation, and back water flooding from Lake Pontchartrain. Nutrient levels were generally low except in the immediate vicinity of the outfall. Mean net primary production of the freshwater forest system was significantly higher downstream of the effluent discharge (1202 g m⁻² yr⁻¹) compared to the control site (799 g m⁻² yr⁻¹). Downstream of the outfall, accretion rates were double the rate of relative sea level rise in the area. Removal efficiencies of N and P were as high as 75% and 95%, respectively. The relatively constant flow of secondarily treated municipal effluent buffered the downstream area from salinity intrusion during a region-wide drought. Re-direction of nutrient-enhanced effluents from open water bodies to wetland ecosystems can maintain plant productivity, sequester carbon, and maintain coastal wetland elevations in response to sea-level rise in addition to improving overall surface water quality, reducing energy use, and increasing financial savings.     

Dynamics of nitrogen and phosphorus retention during wetland ecosystem succession

We compared the mechanisms of nitrogen (N) and phosphorus (P) removal in four young (<15 years old) constructed estuarine marshes with paired mature natural marshes to determine how nutrient retention changes during wetland ecosystem succession. In constructed wetlands, N retention begins as soon as emergent vegetation becomes established and soil organic matter starts to accumulate, which is usually within the first 1–3 years. Accumulation of organic carbon in the soil sets the stage for denitrification which, after 5–10 years, removes approximately the same amount of N as accumulating organic matter, 5–10 g/m²/yr each, under conditions of low N loadings. Under high N loadings, the amount of N stored in accumulating organic matter doubles while N removal from denitrification may increase by an order of magnitude or more. Both organic N accumulation and denitrification provide for long-term reliable N removal regardless of N loading rates. Phosphorus removal, on the other hand, is greatest during the first 1–3 years of succession when sediment deposition and sorption/precipitation of P are greatest. During this time, constructed marshes may retain from 3 g P/m²/yr under low P loadings to as much as 30 g P/m²/yr under high loadings. However, as sedimentation decreases and sorption sites become saturated, P retention decreases to levels supported by organic P accumulation (1–2 g P/m²/yr) and sorption/precipitation with incoming aqueous and particulate Fe, Al and Ca. Phosphorus cycling in wetlands differs from forest and other terrestrial ecosystems in that conservation of P is greatest during the early years of succession, not during the middle or late stages. Conservation of P by wetlands is largely regulated by geochemical processes (sorption, precipitation) which operate independently of succession. In contrast, the conservation of N is controlled by biological processes (organic matter accumulation, denitrification) that change as succession proceeds.     

The 1994 experimental opening of the Bonnet Carre Spillway to divert Mississippi River water into Lake Pontchartrain, Louisiana

A diversion of Mississippi River water into Lake Pontchartrain, Louisiana, USA by way of the Bonnet Carre Spillway has been proposed as a restoration technique to help offset regional wetland loss. An experimental diversion of Mississippi River water into Lake Pontchartrain was carried out in April 1994 to monitor the fate of nutrients and sediments in the spillway and Lake Pontchartrain. Approximately 6.4×10⁸ m³ of Mississippi River water was diverted into Lake Pontchartrain over 42 days. As water passed through the Bonnet Carre Spillway, there were reductions in total suspended sediment concentrations of 82–83%, nitrite+nitrate (NO_x) of 28–42%, in total nitrogen (TN) of 26–30%, and in total phosphorus (TP) of 50–59%. 3.9±1.1 cm of accretion was measured in the spillway. Nutrient concentrations at the freshwater plume edge in Lake Pontchartrain compared to the Mississippi River were lower for NO_x (44–81%), TN (37–57%), and TP (40–70%), and generally higher for organic nitrogen (−7–57%). The Si:N ratio generally increased and the N:P ratio decreased from the river to the plume edge. Nutrient stoichiometric ratios indicate water at the plume edge was not silicate limited, suggesting conditions favoring diatomic phytoplankton.     

Distribution and release of sedimentary phosphorus in Lake Ladoga

Sedimentary phosphorus fractions and phosphorus release from the sediments were studied in Lake Ladoga at altogether 46 sampling sites, representing the full range of sediment types encountered in the lake. Determination of P fractions and physico-chemical analyses were made of surface sediment cores (10–20 cm long, each sampled at 3–4 levels) and in the overlying water. The range of total phosphorus per dry weight of sediment was 0.2–3.3 mg g^–1, and that of inorganic P 0.1–2.5 mg g^–1. The levels of interstitial soluble phosphorus, range 2–613 µg 1^–1 for total P and 1–315 µg 1^–1 for inorganic P, were higher than those of dissolved P concentrations in the overlying water. Diffusive fluxes of phosphate from sediment to the overlying water were estimated using three independent methods. The estimated range was 4–914 µg P m^–2 d^–1; the mean value for the whole bottom area, 0.1 mg P m^–2 d^–1, is lower than previously published estimates. The estimated annual contribution of sedimentary inorganic P flux to Lake Ladoga water is equal to 620 tons of P per year, which amounts to more than 10% of the estimated external P load into the lake. 68% of the total diffusive flux emanates from deep water sediments, which are not exposed to seasonal variation of conditions. In deep lakes, such as Lake Ladoga, phosphorus release from the sediments is controlled primarily by diffusive mechanisms. Wave action and currents as well as bioturbation are probably of importance mainly in shallow near-shore areas. Phosphorus release by gas ebullition and macrophytes is considered negligible.     

The influence of seasonal light variations on the growth of Sphaerotilus natans

A 1979 study of the macrobenthos of Ross Barnett Reservoir, Mississippi, identified 20 genera of invertebrates. Benthic invertebrate productivity ranged from 1.45 g m^–2 yr^–1 in profundal zones to 5.07 g m^–2 yr^–1 in littoral zones with major contributors including Hexagenia bilineata, Chaoborus punctipennis, Chironomus attenuatus, Tanypus stellatus and Coelotanypus tricolor. Numerical and species variation were influenced by depth zones, factors associated with reservoir age (17 years), and sediment and vegetative development in the reservoir benthic zones. Littoral productivity, although limited by lack of zone development, was greater than that of flood control reservoirs in the south central United States because of water level stability. Profundal productivity was limited by hypolimnional oxygen stress. Benthos was a major food source of freshwater drum (Aplodinotus grunniens) which forms a significant portion of the reservoir fisheries.     

Retention of soluble organic nutrients by a forested ecosystem

We document an example of a forested watershed at the Coweeta HydrologicLaboratory with an extraordinary tendency to retain dissolved organic matter(DOM) generated in large quantities within the ecosystem. Our objectives weretodetermine fluxes of dissolved organic C, N, and P (DOC, DON, DOP,respectively),in water draining through each stratum of the ecosystem and synthesizeinformation on the physicochemical, biological and hydrologic factors leadingtoretention of dissolved organic nutrients in this ecosystem. The ecosystemretained 99.3, 97.3, and 99.0% of water soluble organic C, N and P,respectively, produced in litterfall, throughfall, and root exudates. Exportsinstreamwater were 4.1 kg ha^–1yr^–1of DOC, 0.191 kg ha^–1 yr^–1 ofDON, and 0.011 kg ha^–1 yr^–1 ofDOP. Fluxes of DON were greater than those of inorganic N in all strata. MostDOC, DON, and DOP was removed from solution in the A and B horizons, with DOCbeing rapidly adsorbed to Fe and Al oxyhydroxides, most likely by ligandexchange. DON and DOC were released gradually from the forest floor over theyear. Water soluble organic C produced in litterfall and throughfall had adisjoint distribution of half-decay times with very labile and veryrefractory fractions so that most labile DOC was decomposed before beingleachedinto the mineral soil and refractory fractions dominated the DOC transportedthrough the ecosystem. We hypothesize that this watershed retained solubleorganic nutrients to an extraordinary degree because the soils have very highcontents of Fe and Al oxyhydroxides with high adsorption capacities and becausethe predominant hydrologic pathway is downwards as unsaturated flow through astrongly adsorbing A and B horizon. The well recognized retention mechanismsforinorganic nutrients combine with adsorption of DOM and hydrologic pathway toefficiently prevent leaching of both soluble inorganic andorganic nutrients in this watershed.     

Sediment nutrient accumulation and nutrient availability in two tidal freshwater marshes along the Mattaponi River, Virginia, USA

Sediment deposition is the main mechanism of nutrient delivery to tidal freshwater marshes (TFMs). We quantified sediment nutrient accumulation in TFMs upstream and downstream of a proposed water withdrawal project on the Mattaponi River, Virginia. Our goal was to assess nutrient availability by comparing relative rates of carbon (C), nitrogen (N), and phosphorus (P) accumulated in sediments with the C, N, and P stoichiometries of surface soils and above ground plant tissues. Surface soil nutrient contents (0.60–0.92% N and 0.09–0.13% P) were low but within reported ranges for TFMs in the eastern US. In both marshes, soil nutrient pools and C, N, and P stoichiometries were closely associated with sedimentation patterns. Differences between marshes were more striking than spatial variations within marshes: both C, N, and P accumulation during summer, and annual P accumulation rates (0.16 and 0.04 g P m^–2 year^–1, respectively) in sediments were significantly higher at the downstream than at the upstream marsh. Nitrogen:P ratios <14 in above ground biomass, surface soils, and sediments suggest that N limits primary production in these marshes, but experimental additions of N and/or P did not significantly increase above ground productivity in either marsh. Lower soil N:P ratios are consistent with higher rates of sediment P accumulation at the downstream site, perhaps due to its greater proximity to the estuarine turbidity maximum.     

Gull contributions of phosphorus and nitrogen to a Cape Cod kettle pond

Nutrient excretion rates and the annual contribution of P from the feces of the gulls Larus argentatus and L. marinus (and of N from L. argentatus) to the nutrient budget of Gull Pond (Wellfleet), a soft water seepage lake, have been estimated. Intensive year-round gull counts by species were combined with determinations of defecation rate and the nutrient content of feces to quantitatively assess the P loading rates associated with regular gull use of this coastal pond on a seasonal and annual basis. Total P loading from gulls was estimated to be 52 kg yr^–1, with 17 kg from L. argentatus and 35 kg from L. marinus, resulting from about 5.0 × 10⁶ h yr^–1 and 1.7 × 10⁶ h yr^–1 of pond use. This compares with P loading estimates of 67 kg yr^–1 from upgradient septic systems, 2 kg yr^–1 from precipitation and 3 kg yr^–1 from unpolluted ground water. Fifty-six percent of annual gull P loading was associated with migratory activity in late fall. Estimated annual N loading by L. argentatus was 14 kg TKN, 206 g NO₃-N, and 1.85 g g NH₃-N.     

Freshwater marshes as dissolved silica recyclers in an estuarine environment (Schelde estuary, Belgium)

Compared to knowledge about N and P processing in the aquatic continuum of lakes, wetlands and estuaries, knowledge concerning transport and cycling of Si is only fragmentary. Furthermore, Si research in estuaries has mainly been focused on subtidal benthic sediments and uptake and recycling by diatom communities. The biogeochemical cycling of Si in tidal wetlands, which can contain large amounts of Si, has thus far been neglected. We have conducted several whole ecosystem Si mass-balances on a freshwater marsh located in the Schelde estuary (6 tidal cycles, 2 with BSi included). Our measurements show that the freshwater marsh acts as an important source of dissolved Si to the main river (1–18% more export than import, on average 0.114 g m^–2). This export is compensated by import of amorphous silica into the marsh (19–55% more import than export). The marsh was shown to act as silica recycler, resupplying biologically available dissolved Si to the estuarine ecosystem. Extrapolations show that during summer and spring months, when dissolved silica is depleted due to diatom growth, almost half of the total dissolved silica load in the main river channel could result from marsh recycling.