Organic carbon dynamics in two regulated rivers in northwestern Montana,USA

We studied the transport of particulate organic carbon (POC) and dissolved organic carbon (DOC) in two regulated rivers during minimum and increasing discharges. Mean annual concentrations of total POC, measured monthly during conditions of minimum discharge from the dams, were twice as high at a station below a dam with a selective withdrawal system on the Kootenai River (KR, 0.15 mg 1^–1), as at station below a dam with hypolimnetic water releases on the Flathead River (FR, 0.07 mg 1^–1). Annual mean concentrations of DOC were similar below both dams (1.62 mg 1^–1 FR; 1.71 KR). The percentage of POC in four size fractions differed in regulated and unregulated reaches of each river system; the smallest size fraction (0.45–10 smm) constituted a larger percentage of the total POC at the stations below the dams (50–93%), because POC in large size classes had settled out in the reservoir. The three largest size fractions (10–1000 µm) comprised a larger percentage of the total POC when samples were taken during conditions of full discharge from the dam. We measured large increases in all size classes of POC in samples collected during increasing discharges in a regulated reach, reflecting the component of sloughed periphyton and resuspended organic matter that were added during periods of hydropower generation at the dam. Seston (355 µm to 1 cm) collected in nets increased dramatically during increasing flows; concentrations of particulate organic matter (POM) in samples collected two and three hours after water levels began to rise were 572 and 1440 times higher than those collected during minimum discharge at the dam.     

Organic contaminant exposure in the Lake St. Clair food web

Chemical concentrations and distributions in an aquatic food web were studied to quantify the relative importance of chemical properties versus food web processes in determining exposure dynamics of organic contaminants in aquatic ecosystems. Five organochlorines were measured (Pentachlorobenzene QCB, Hexachlorobenzene HCB, Octachlorostyrene OCS, Dichlorodiphenyldichloroethylene DDE and Polychlorinated Biphenyls PCBs) in the food web of Lake St. Clair. Levels of QCB in aquatic organisms ranged from 1.0 to 25 µg kg^–1 lipid, and levels of HCB ranged from 10 to 410 µg kg^–1 lipid. More elevated concentrations of OCS (13 to 392 µg kg^–1 lipid), DDE (162 to 11 986 µg kg^–1 lipid) and PCB (650 to 64 900 µg kg^–1 lipid) were observed. Organism — water equilibrium ratios were calculated for all species sampled to quantify the importance of food web processes in regulating contaminant exposure dynamics. Correlations of organism — water equilibrium ratios with body size were not significant for QCB, HCB and OCS (P>0.1), but were found to be significant for DDE and PCB (P<0.01).Results support the conclusion that both chemical properties and food web dynamics regulate the distribution and concentration of organochlorines in aquatic ecosystems. Food web processes are important, however, for chemicals, that are not metabolized and have octanol — water partition coefficients (log K _ow) greater than 5.5.     

Impacts of Reservoir Creation on the Biogeochemical Cycling of Methyl Mercury and Total Mercury in Boreal Upland Forests

The FLooded Upland Dynamics Experiment (FLUDEX) at the Experimental Lakes Area (ELA) in northwest Ontario was designed to test the hypothesis that methylmercury (MeHg) production in reservoirs is related to the amount, and subsequent decomposition, of flooded organic matter. Three upland forest sites that varied in the amounts of organic carbon stored in vegetation and soils (Low C, 30,870 kg C ha⁻¹; Medium C, 34,930 kg C ha⁻¹; and High C, 45,860 kg C ha⁻¹) were flooded annually from May to September with low-organic carbon, low-MeHg water pumped from a nearby lake. Within five weeks of flooding, MeHg concentrations in the reservoir outflows exceeded those in reservoir inflows and remained elevated for the duration of the experiment, peaking at 1.60 ng L⁻¹ in the Medium C reservoir. We estimated the net production of MeHg in each reservoir by calculating annual changes in pools of MeHg stored in flooded soils, periphyton, zooplankton, and fish. Overall, there was an initial pulse of MeHg production (range = 120–1590 ng m⁻² day⁻¹) in all FLUDEX reservoirs that lasted for 2 years, after which time net demethylation (range = 360–1230 ng MeHg degraded m⁻² day⁻¹) began to reduce the pools of MeHg in the reservoirs, but not back to levels found prior to flooding. Rates of MeHg production were generally related to the total amount of organic carbon flooded to create the reservoirs. Large increases in MeHg stores in soils compared to those in water and biota indicate that flooded soils were the main sites of MeHg production. This study should assist hydroelectric utilities and government agencies in making informed decisions about selecting sites for future reservoir development to reduce MeHg contamination of the reservoir fisheries.     

Carbon Dioxide and Methane Production in Small Reservoirs Flooding Upland Boreal Forest

The FLooded Uplands Dynamics EXperiment (FLUDEX) was designed to assess the impact of reservoir creation on carbon cycling in boreal forests by (a) determining whether production of the greenhouse gases (GHG) carbon dioxide (CO₂) and methane (CH₄) in reservoirs is related to the amount of organic carbon (OC) stored in the flooded landscape, (b) examining temporal trends in GHG production during initial stages of flooding, and (c) considering the net difference between GHG fluxes before and after flooding to estimate the true effect of reservoir creation on atmospheric GHG levels. Three forested sites that varied in the amount of OC stored in soils and vegetation (30,870–45,860 kg C ha^–1) were experimentally flooded from June to September in 1999–2001. Throughout the study, net CO₂ and CH₄ production in all three reservoirs was not related to overall site OC storage. During the 1st flooding season, net CO₂ production in the three reservoirs was 703–797 kg C ha^–1, but it decreased during the 2nd and 3rd flooding seasons to between 408 and 479 kg C ha^–1. However, CH₄ production increased in all reservoirs with each flooding season, from about 3.2–4.6 kg C ha^–1 in 1999 to 12.8–24.9 kg C ha^–1 in 2000 and 29.7–35.2 kg C ha^–1 in 2001. Over the long term, effects of boreal reservoir creation on atmospheric GHG levels may be largely due to net changes in CH₄ cycling between the undisturbed and flooded ecosystems.     

Distribution and Physiological Characteristics of Hyperthermophiles in the Kubiki Oil Reservoir in Niigata, Japan

The distribution of culturable hyperthermophiles was studied in relation to environmental conditions in the Kubiki oil reservoir in Japan, where the temperature was between 50 and 58°C. Dominant hyperthermophilic cocci and rods were isolated and shown to belong to the genera Thermococcus and Thermotoga, respectively, by 16S rDNA analyses. Using the most-probable-number method, we found that hyperthermophilic cocci were widely distributed in several unconnected fault blocks in the Kubiki oil reservoir. In 1996 to 1997, their populations in the production waters from oil wells were 9.2 × 10³ to 4.6 × 10⁴ cells/ml, or 10 to 42% of total cocci. On the other hand, hyperthermophilic rods were found in only one fault block of the reservoir with populations less than 10 cells/ml. Dominant Thermococcus and Thermotoga spp. grew at reservoir temperatures and utilized amino acids and sugars, respectively, as sole carbon sources. While organic carbon was plentiful in the environment, these hyperthermophiles were unable to grow in the formation water due to lack of essential nutrients. Concentrations of some organic and inorganic substances differed among fault blocks, indicating that the movement of formation water between fault blocks was restricted. This finding suggests that the supply of nutrients via fluid current is limited in this subterranean environment and that the organisms are starved in the oil reservoir. Under starved conditions at 50°C, culturable cells of Thermococcus sp. remained around the initial cell density for about 200 days, while those of Thermotoga sp. decreased exponentially to 0.01% of the initial cell density after incubation for the same period. The difference in survivability between these two hyperthermophiles seems to reflect their populations in the fault blocks. These results indicate that hyperthermophilic cocci and rods adapt to the subterranean environment of the Kubiki oil reservoir by developing an ability to survive under starved conditions.     

Transport and release of chemicals from plastics to the environment and to wildlife

Plastics debris in the marine environment, including resin pellets, fragments and microscopic plastic fragments, contain organic contaminants, including polychlorinated biphenyls (PCBs), polycyclic aromatic hydrocarbons, petroleum hydrocarbons, organochlorine pesticides (2,2′-bis(p-chlorophenyl)-1,1,1-trichloroethane, hexachlorinated hexanes), polybrominated diphenylethers, alkylphenols and bisphenol A, at concentrations from sub ng g^–1 to µg g^–1. Some of these compounds are added during plastics manufacture, while others adsorb from the surrounding seawater. Concentrations of hydrophobic contaminants adsorbed on plastics showed distinct spatial variations reflecting global pollution patterns. Model calculations and experimental observations consistently show that polyethylene accumulates more organic contaminants than other plastics such as polypropylene and polyvinyl chloride. Both a mathematical model using equilibrium partitioning and experimental data have demonstrated the transfer of contaminants from plastic to organisms. A feeding experiment indicated that PCBs could transfer from contaminated plastics to streaked shearwater chicks. Plasticizers, other plastics additives and constitutional monomers also present potential threats in terrestrial environments because they can leach from waste disposal sites into groundwater and/or surface waters. Leaching and degradation of plasticizers and polymers are complex phenomena dependent on environmental conditions in the landfill and the chemical properties of each additive. Bisphenol A concentrations in leachates from municipal waste disposal sites in tropical Asia ranged from sub µg l^–1 to mg l^–1 and were correlated with the level of economic development.     

Seasonal Patterns of Soil Respiration and Related Soil Biochemical Properties under Nitrogen Addition in Winter Wheat Field

Understanding the changes of soil respiration under increasing N fertilizer in cropland ecosystems is crucial to accurately predicting global warming. This study explored seasonal variations of soil respiration and its controlling biochemical properties under a gradient of Nitrogen addition during two consecutive winter wheat growing seasons (2013–2015). N was applied at four different levels: 0, 120, 180 and 240 kg N ha^-1 year^-1 (denoted as N0, N12, N18 and N24, respectively). Soil respiration exhibited significant seasonal variation and was significantly affected by soil temperature with Q₁₀ ranging from 2.04 to 2.46 and from 1.49 to 1.53 during 2013–2014 and 2014–2015 winter wheat growing season, respectively. Soil moisture had no significant effect on soil respiration during 2013–2014 winter wheat growing season but showed a significant and negative correlation with soil respiration during 2014–2015 winter wheat growing season. Soil respiration under N24 treatment was significantly higher than N0 treatment. Averaged over the two growing seasons, N12, N18 and N24 significantly increased soil respiration by 13.4, 16.4 and 25.4% compared with N0, respectively. N addition also significantly increased easily extractable glomalin-related soil protein (EEG), soil organic carbon (SOC), total N, ammonium N and nitrate N contents. In addition, soil respiration was significantly and positively correlated with β-glucosidase activity, EEG, SOC, total N, ammonium N and nitrate N contents. The results indicated that high N fertilization improved soil chemical properties, but significantly increased soil respiration.     

Effects of Seasonal Upwelling on Inorganic and Organic Matter Dynamics in the Water Column of Eastern Pacific Coral Reefs

The Gulf of Papagayo at the northern Pacific coast of Costa Rica experiences pronounced seasonal changes in water parameters caused by wind-driven coastal upwelling. While remote sensing and open water sampling already described the physical nature of this upwelling, the spatial and temporal effects on key parameters and processes in the water column have not been investigated yet, although being highly relevant for coral reef functioning. The present study investigated a range of water parameters on two coral reefs with different exposure to upwelling (Matapalo and Bajo Rojo) in a weekly to monthly resolution over one year (May 2013 to April 2014). Based on air temperature, wind speed and water temperature, three time clusters were defined: a) May to November 2013 without upwelling, b) December 2013 to April 2014 with moderate upwelling, punctuated by c) extreme upwelling events in February, March and April 2014. During upwelling peaks, water temperatures decreased by 7°C (Matapalo) and 9°C (Bajo Rojo) to minima of 20.1 and 15.3°C respectively, while phosphate, ammonia and nitrate concentrations increased 3 to 15-fold to maxima of 1.3 μmol PO₄ ^3- L^-1, 3.0 μmol NH₄ ⁺ L^-1 and 9.7 μmol NO₃ ^- L^-1. This increased availability of nutrients triggered several successive phytoplankton blooms as indicated by 3- (Matapalo) and 6-fold (Bajo Rojo) increases in chlorophyll a concentrations. Particulate organic carbon and nitrogen (POC and PON) increased by 40 and 70% respectively from February to April 2014. Dissolved organic carbon (DOC) increased by 70% in December and stayed elevated for at least 4 months, indicating high organic matter release by primary producers. Such strong cascading effects of upwelling on organic matter dynamics on coral reefs have not been reported previously, although likely impacting many reefs in comparable upwelling systems.     

Physicochemical limnology of the Tongue River Reservoir,Montana

A one year physicochemical survey was conducted on the Tongue River Reservoir, a run of the river impoundment in southeastern Montana. The Tongue River was the only significant inflow and supplied 93, 96 and 97% of the nutrient, major ion and water inputs to the impoundment. Heat advected from inflowing water accounted for 17% of the energy gained during the summer heating cycle. The annual nutrient load to the reservoir from the river was 20.2 g m^–2 total nitrogen (TN) and 3.8 g m^–2 total phosphorus (TP). Due to the absence of reducing conditions at depth and the complex seasonal pattern of water movement through the reservoir, 99% of the TN load was discharged but 49% of the TP load was retained in the reservoir.     

The use of carbon isotope ratios to evaluate legume contribution to soil enhancement in tropical pastures

Soil carbon distribution with depth, stable carbon isotope ratios in soil organic matter and their changes as a consequence of the presence of legume were studied in three 12-year-old tropical pastures (grass alone —Brachiaria decumbens (C₄), legume alone —Pueraria phaseoloides (C₃) and grass + legume) on an Oxisol in Colombia. The objective of this study was to determine the changes that occurred in the¹³C isotope composition of soil from a grass + legume pasture that was established by cultivation of a native savanna dominated by C₄ vegetation. The¹³C natural abundance technique was used to estimate the amount of soil organic carbon originating from the legume. Up to 29% of the organic carbon in soil of the grass + legume pasture was estimated to be derived from legume residues in the top 0–2-cm soil depth, which decreased to 7% at 8–10 cm depth. Improvements in soil fertility resulting from the soil organic carbon originated from legume residues were measured as increased potential rates of nitrogen mineralization and increased yields of rice in a subsequent crop after the grass + legume pasture compared with the grass-only pasture. We conclude that the¹³C natural abundance technique may help to predict the improvements in soil quality in terms of fertility resulting from the presence of a forage legume (C₃) in a predominantly C₄ grass pasture.     

Stability of sodium and potassium complexes of valinomycin

Stability constants of sodium and potassium complexes of valinomycin in some alcohols and water—organic solvent mixtures have been determined by titration, using circular dichroism to monitor complex formation. Constants range from 10¹ to 10⁶ M⁻¹. Stability of the potassium and sodium complexes increases with decreasing dielectric constant, but the ratio of the constants remains about 10³–10⁴. As others have shown, a similar selectivity for K⁺ is observed in a number of other types of measurements involving valinomycin. These include the permeability and conductance ratios which characterize the selectivity of cation transport through membranes and the ratio of salt extraction equilibrium constants. On the basis of data presented here, and elsewhere, it is suggested that conformational constraints within the depsipeptide part of the complexes aid ion selectivity and that differences in cation solvation and carbonyl ligand binding energies make an important, roughly equal, contribution.     

Food consumption by the larvae of Sericostoma vittatum (Trichoptera), an endemic species from the Iberian Peninsula

The caddisfly Sericostoma vittatum Rambur (Trichoptera: Sericostomatidae) is an endemic species of the Iberian Peninsula. Under laboratory conditions, larvae of S. vittatum had a higher activity and metabolism during the night. Besides consuming particulate allochthonous organic matter, young stages are also able to feed and grow on faecal pellets from adults. Daily growth rates varied from 0.02 mg (0.8–3.7 mg size class animals) to 0.31 mg dry mass (10.6–22.8 mg size class animals). Due to the high densities of this species (annual mean of 25 individuals m^–2; maximum of 96 individuals m^–2) and high consumption rates (0.47 mg leaf dry mass mg animal^–1 d^–1for small larvae), this species has a potential key role on the fragmentation of allochthonous organic matter of streams in central Portugal.     

The genetic map of the mitochondrial genome in yeast

Summary An approach for the screening of mit ^- mutants, the isolation and preliminary classification of a series of such mutants is reported. Loss and retention of 8 mit ^- and 6 drug ^r markers in mitDNA was analyzed in populations of rho^- clones derived from four yeast strains. The populations studied constitute a representative fraction of the rho^- petites formed during growth at 35° C under the influence of mutation tsp-25 which is in common to the four strains. The majority of the rho^- clones retained several of the markers studied. Depending on the marker regarded retention frequencies between 15% (oxi3) and 45% (oli1, cob) were observed. Loss of one and retention of the other of a pair of markers was determined in all rho^- clones of the four populations. The frequencies of marker separation by rho^- deletion thus obtained are assumed to reflect the distance between markers on the mitochondrial genome: the higher the frequency of separation the longer the distance between two markers. Based on these frequencies a unique order of markers on a circular map was determined. Positions of markers on a scale from 0 to 100 were found to be: cap/ery (0) — olil (16) — cob1-1354 (21) — ana101 (22) — cob2-1625 (24) — oli2 (35) — pho1 (40) — oxi3-2501 (44) — oxi3-3771 (47) — par (65) — oxi2 (79) — oxil (87) tms8 (93) —cap (100). The relevance of this map as to the faithful representation of the topology of gene loci on mitDNA is discussed. Correlation of retention frequencies of markers to their map positions reveals a pronounced polarity: mitDNA segments carrying the cob-oli1 segment prevail whereas segments retaining oxi3 are the least frequent.     

Phosphorus release from resuspended sediment in the shallow and wind-exposed Lake Arresø, Denmark

Wind-induced sediment resuspension occurs frequently in the shallow and eutrophic Lake Arresø, Denmark. The impact of resuspension on internal phosphorus loading was investigated by laboratory experiments studying P-release from the undisturbed sediment surface and by experiments simulating resuspension events.Phosphorus release from undisturbed sediment sampled in May and August was 12 mg and 4 mg m^–2 d^–1, respectively. During experimental simulation of resuspension, soluble reactive phosphate (SRP) increased by 20–80 µg l^–1, which indicates that a typical resuspension event in the lake would be accompanied by the release of 150 mg SRP m^–2. The internal P loading induced by resuspension is estimated to be 60–70 mg m^–2 d^–1, or 20–30 times greater than the release from undisturbed sediment.SRP release during simulation of resuspension was mainly dependent on the equilibrium conditions in the water column and was basically independent of the increase in suspended solids and the duration of resuspension. A second simulation of resuspension conducted 26 hours later, did not result in any further release of SRP from sediment sampled in May. In contrast, there was an additional SRP release from sediment sampled in August, indicating that an exchangable P pool, capable of altering equilibrium conditions, is built up between resuspension events.It is concluded that resuspension, by increasing the P flux between sediment and water, plays a major role in the maintenance of the high nutrient level in Lake Arresø. A relatively high release rate is maintained during resuspension because of the low Fe:P ratio and the high concentration of NH₄Cl-extractable P in the sediment.     

Phase diagram of lipid A from Salmonella minnesota and Escherichia coli rough mutant lipopolysaccharide

We have reported here on the structural polymorphism of lipid A, the “endotoxic principle” of bacterial lipopolysaccharide. For lipid A of rough mutant lipopolysaccharide from Salmonella minnesota and Escherichia coli, the three-dimensional supramolecular structures were determined with x-ray diffraction utilizing synchrotron radiation. The investigations were performed in the water concentration range 10 to 95% by weight, at [lipid A]:[Mg²⁺] molar ratios from 1:0 to 0.1:1, and in the temperature range from 20 to 70°C. These data were correlated with measurements of the β→α phase behaviour which was monitored with differential scanning calorimetry and Fourier-transform infrared spectroscopy. We found that the transition temperature of the acyl chains ranges—in the absence of Mg²⁺—from 45°C at high to 56°C at low water content, and—at an equimolar content of Mg²⁺—from 52°C at high to 59°C at low water concentrations. In the gel phase—in which the lipid A acyl chains are more disordered than those from saturated phospholipids—cubic phases are adopted at high water content (>60%) and at high [lipid A):[Mg²⁺] molar ratios. At low water contents, lamellar states are assumed exclusively. In the liquid crystalline state of lipid A, the hexagonal H_II, state is adopted under all conditions. The structural variability of lipid A is highest at high water concentrations, and structural changes may be induced by only slight changes in temperature, water content, and Mg²⁺ concentration. Under physiological conditions, however, the lipid A assemblies exhibit a strong preference to cubic structures.     

Chemical limnology of soft water lakes in the Upper Midwest

Water samples from 36 lakes in northern Minnesota, Wisconsin, and Michigan were collected and analyzed during 1983–1984. All study lakes were dilute and had total alkalinities of less than 150 eq · L^–1. Minnesota lakes have hydrologic inputs from the watershed and inputs of base cations derived from the watershed. Study lakes in Minnesota had higher total alkalinities, dissolved organic carbon, and noncarbonate alkalinity as a result of watershed inputs. Lakes in Michigan and Wisconsin were precipitation-dominated seepage lakes that have lower concentrations of base cations than lakes in Minnesota. All of the study lakes have lower sulfate concentrations than expected, based on atmospheric wet deposition and evapotranspiration.Pore water samples collected from one of the study lakes—Little Rock Lake—in Wisconsin were used to calculate diffusive fluxes between the sediment and water column. According to these calculations, the sediments were a source of total alkalinity and Ca²⁺ and a sink for SO₄ ^2–. The sediment-water exchange of total alkalinity, Ca²⁺, and SO₄ ^2– appears to be important in the whole-lake budgets of these ions for Little Rock Lake.     

Dynamics of transparent exopolymeric particles (TEP) production by Phaeocystis globosa under N- or P-limitation: a controlling factor of the retention/export balance

The concentration of transparent exopolymeric particles (TEP) was monitored during Phaeocystis globosa blooms that developed in mesocosms under different initial N:P ratios (from N- to P-limited conditions). TEP concentration was measured using the microscopic (TEP_micro, ppm) and the colorimetric (TEP_color, Xanthan equiv. L⁻¹) methods. TEP concentrations varied from 5 to >75 ppm and from 60 to >1500 μg Xanthan equiv. L⁻¹, and were relatively low until the mesocosms reached nutrient (either N or P) depletion and then increased abruptly. From the TEP_micro versus TEP_color concentrations comparison and from their relation to chlorophyll a concentrations, two phases for the dynamics of TEP production were identified: (1) production through active release of precursors during the growth phase of P. globosa — defined as TEP₁ — and their integration into the TEP pool through coagulation processes; (2) release of large TEP from the mucilaginous matrix of P. globosa colonies subsequent to disruption caused by nutrient depletion — defined as TEP₂ — and their direct integration into the TEP pool outside the constraint of coagulation. The formation of a multiorigin TEP pool during P. globosa blooms may have implications for the fate of the blooms, due to difference in TEP bioreactivity according to their source and to difference in timing and intensity of TEP₁ versus TEP₂ production according to N- or P-depletion. For P. globosa blooms developing under N-limiting conditions, the transition from the first source (i.e. TEP₁) to the second one (i.e. TEP₂) was a slow and continuous process. In contrast, the P. globosa bloom developing under P-limiting conditions showed the sudden formation of heavy mucous aggregates when P became depleted, that may have been caused by a massive release of TEP₂. Our study suggests that the nutrient regime may control the export vs. retention balance during P. globosa blooms, via production of a multiorigin TEP pool.     

Nitrogen fluxes and budget seasonality in the Ria Vigo [2pt] (NW Iberian Peninsula)

Inorganic and organic nitrogen fluxes in the Ria Vigo have been quantified in order to recognise the contrasting nitrogen budget scenarios and understand the biogeochemical response to eutrophication events. According to the nitrogen biogeochemical pathways of the ria reservoir (photosynthesis, remineralization, denitrification, PON rain rate and sedimentation), three main seasonal behavioural trends are emphasised: (1) low inorganic nitrogen inputs and low organic nitrogen fluxes, (2) high inorganic nitrogen input and output, (3) high inorganic nitrogen input and high organic nitrogen output. The first scenario occurs in late spring and in summer during non-upwelling situations. The consumption of inorganic nitrogen by net photosynthesis is approximately 2 mol N s^–1 and the ria is oligotrophic (12 mgC m^–2 h^–1). The outgoing estuarine residual current transports phytoplanktonic material towards the mouth of the ria whereupon it sediments and is remineralized as it falls to the lower water layers and the incoming residual current. The regenerated nitrogen is reintroduced to the photic ria layer which leads to the greatest reduction in dissolved oxygen concentration (50% of saturation). Recycled nutrients play an important role in primary production during this oligotrophic state of the ria. Thus, approximately half of the inorganic nitrogen utilised by photosynthesis is ammonium. The majority of PON is deposited inside the ria (0.8 mmol N m^–2 d^–1) and the denitrification rate is 0.3 mmol N₂ m^–2 d^–1. The other two cases occur in winter and spring–summer with upwelling. In winter, estuarine circulation and freshwater contributions control the nitrogen cycle. The ria mainly exports nitrate (up to 14 mol N s^–1) and so there is fertilisation but no eutrophication. In spring and summer, the nitrogen cycle is controlled by upwelling circulation. The inorganic nitrogen consumption by net photosynthesis is high, 7–14 mmol N m^–2 d^–1, and the ria is a natural eutrophic system (70 mgC m^–2 h^–1). Accordingly, 90% of organic nitrogen is synthesised from nitrate and the upwelling-increased circulation exports 6.5 mol N s^–1 of organic nitrogen.     

A historic perspective on Wadden Sea eutrophication

In this paper, a reconstruction of the pre-industrial trophic status of the Wadden Sea is presented. A conceptual model is outlined that links the organic matter and nutrient dynamics in the Wadden Sea with riverine nutrient input. Fundamental processes in this model are: a nutrient-limited offshore primary production and the subsequent import of primary produced organic matter from the North Sea into the Wadden Sea. Two approaches have been followed to estimate the production and remineralisation levels under pre-industrial conditions. The first approach is based on present-day relationships between the seasonal cycle of NH₄ and NO₂ in the western Dutch Wadden Sea and suggests, on average, sixfold lower production and remineralisation rates under pre-industrial conditions (range: four to eight times). The second approach is based on present carbon budgets extrapolated to pre-industrial budgets on the basis of present relationships between winter nutrient concentrations, annual primary production and annual organic matter turnover rates, and suggests a fivefold lower organic matter turnover under pre-industrial conditions (annual primary production: ~55 g C m^–2 year^–1, annual remineralisation: ~77 g C m^–2 year^–1). Better pre-industrial light conditions in the Wadden Sea may have allowed a more efficient use of nutrients, a higher annual primary production of about 86 g C m^–2 year^–1 and annual remineralisation rates of about 108 g C m^–2 year^–1.     

Primary production of ice algae on a seasonally-ice-covered,continental shelf

Summary Ice algae samples were collected from the winter pack ice off Labrador during March 1984. The population was dominated by centric diatoms. Chlorophyll concentrations ranged from 40 to 190 mg m^–3, and particulate organic carbon from 2 to 10 g m^–3. Assimilation numbers for the ice algae ranged from 1.4 to 2.8 mg C (mg chl)^–1 h^–1 with a mean of 2.3, and were not significantly different from the assimilation numbers of the pelagic community beneath the ice. The ice algae were not photoinhibited at light intensities approaching surface light intensities. It is postulated that the dynamic nature of the ice field permits near-surface light intensities to reach the ice algae community at irregular intervals thereby suppressing photoinhibition.