The Status and Characteristics of Eutrophication in the Yangtze River (Changjiang) Estuary and the Adjacent East China Sea, China

Eutrophication has become increasingly serious and noxious algal blooms have been of more frequent occurrence in the Yangtze River Estuary and in the adjacent East China Sea. In 2003 and 2004, four cruises were undertaken in three zones in the estuary and in the adjacent sea to investigate nitrate (NO₃–N), ammonium (NH₄–N), nitrite (NO₂–N), soluble reactive phosphorus (SRP), dissolved reactive silica (DRSi), dissolved oxygen (DO), phytoplankton chlorophyll a (Chl a) and suspended particulate matter (SPM). The highest concentrations of DIN (NO₃–N+NH₄–N+NO₂–N), SRP and DRSi were 131.6, 1.2 and 155.6 μM, respectively. The maximum Chl a concentration was 19.5 mg m⁻³ in spring. An analysis of historical and recent data revealed that in the last 40 years, nitrate and SRP concentrations increased from 11 to 97 μM and from 0.4 to 0.95 μM, respectively. From 1963 to 2004, N:P ratios also increased from 30–40 up to 150. In parallel with the N and P enrichment, a significant increase of Chl a was detected, Chl a maximum being 20 mg m⁻³, nearly four times higher than in the 1980s. In 2004, the mean DO concentration in bottom waters was 4.35 mg l⁻¹, much lower than in the 1980s. In comparison with other estuaries, the Yangtze River Estuary was characterized by high DIN and DRSi concentrations, with low SRP concentrations. Despite the higher nutrient concentrations, Chl a concentrations were lower in the inner estuary (Zones 1 and 2) than in the adjacent sea (Zone 3). Based on nutrient availability, SPM and hydrodynamics, we assumed that in Zones 1 and 2 phytoplankton growth was suppressed by high turbidity, large tidal amplitude and short residence time. Furthermore, in Zone 3 water stratification was also an important factor that resulted in a greater phytoplankton biomass and lower DO concentrations. Due to hydrodynamics and turbidity, the open sea was unexpectedly more sensitive to nutrient enrichment and related eutrophication processes.     

Effects of zooplankton on nutrient availability and seston C : N : P stoichiometry in inshore waters of Lake Biwa, Japan

Forty-eight-hour experimental manipulations of zooplankton biomass were performed to examine the potential effects of zooplankton on nutrient availability and phytoplankton biomass (as measured by seston concentration) and C : N : P stoichiometry in eutrophic nearshore waters of Lake Biwa, Japan. Increasing zooplankton, both mixed-species communities and Daphnia alone, consistently reduced seston concentration, indicating that nearshore phytoplankton were generally edible. The zooplankton clearance rates of inshore phytoplankton were similar to rates measured previously for offshore phytoplankton. Increased zooplankton biomass led to increased concentrations of nutrients (NH₄-N, soluble reactive phosphorus [SRP]). Net release rates were higher than those found in previous measurements made offshore, reflecting the nutrient-rich nature of inshore seston. Zooplankton nutrient recycling consistently decreased TIN : SRP ratios (TIN = NH₄ + NO₃ + NO₂). This effect probably resulted from the low N : P ratios of nearshore seston, which were lower than those commonly found in crustacean zooplankton and thus resulted in low retention efficiency of P (relative to N) by the zooplankton. Thus, zooplankton grazing inshore may ameliorate algal blooms due to direct consumption but tends to create nutrient supply conditions with low N : P, potentially favoring cyanobacteria. In comparison with previous findings for offshore, it appears that potential zooplankton effects on phytoplankton and nutrient dynamics differ qualitatively in inshore and offshore regions of Lake Biwa. Received: September 4, 2000 / Accepted: January 23, 2001     

Thermal stratification, nutrient dynamics, and phytoplankton productivity during the onset of spring phytoplankton growth in Lake Baikal, Russia

Lake Baikal, Russian Siberia, was sampled in July 1990 during the period of spring mixing and initiation of thermal stratification. Vertical profiles of temperature, dissolved nutrients (nitrate and soluble reactive phosphorus), phytoplankton biomass, and primary productivity were determined in an eleven-station transect encompassing the entire 636 km length of the lake. Pronounced horizontal variability in hydrodynamic conditions was observed, with the southern region of the lake being strongly thermally stratified while the middle and north basins were largely isothermal through July. The extent of depletion of surface water nutrients, and the magnitude of phytoplankton biomass and productivity, were found to be strongly correlated with the degree of thermal stratification. Horizontal differences likely reflected the contribution of two important factors: variation in the timing of ice-out in different parts of the lake (driving large-scale patterns of thermal stratification and other limnological properties) and localized effects of river inflows that may contribute to the preliminary stabilization of the water column in the face of intense turbulent spring mixing (driving meso-scale patterns). Examination of the relationships between surface water inorganic N and P depletion suggested that during the spring and early summer, phytoplankton growth in unstratified portions of the lake was largely unconstrained by nutrient supplies. As summer progressed, the importance of co-limitation by both N and P became more apparent. Uptake and regeneration rates, measured directly using the stable isotope ¹⁵N, revealed that phytoplankton in stratified portions of the lake relied primarily on NH₄ as their N source. Rates of NH₄ regeneration were in approximate equilibrium with uptake; both processes were dominated by organisms <2 µm. This pattern is similar to that observed for oligotrophic marine systems. Our study underscores the importance of hydrodynamic conditions in influencing patterns of biological productivity and nutrient dynamics that occur in Lake Baikal during its brief growing season.     

Is inorganic nutrient enrichment a driving force for the formation of red tides?: A case study of the dinoflagellate Scrippsiella trochoidea in an embayment

Red tides (high biomass phytoplankton blooms) have frequently occurred in Hong Kong waters, but most red tides occurred in waters which are not very eutrophic. For example, Port Shelter, a semi-enclosed bay in the northeast of Hong Kong, is one of hot spots for red tides. Concentrations of ambient inorganic nutrients (e.g. N, P), are not high enough to form the high biomass of chlorophyll a (chl a) in a red tide when chl a is converted to its particulate organic nutrient (N) (which should equal the inorganic nutrient, N). When a red tide of the dinoflagellate Scrippsiella trochoidea occurred in the bay, we found that the red tide patch along the shore had a high cell density of 15,000 cells ml⁻¹, and high chl a (56 μg l⁻¹), and pH reached 8.6 at the surface (8.2 at the bottom), indicating active photosynthesis in situ. Ambient inorganic nutrients (NO₃, PO₄, SiO₄, and NH₄) were all low in the waters and deep waters surrounding the red tide patch, suggesting that the nutrients were not high enough to support the high chl a >50 μg l⁻¹ in the red tide. Nutrient addition experiments showed that the addition of all of the inorganic nutrients to a non-red-tide water sample containing low concentrations of Scrippsiella trochoidea did not produce cell density of Scrippsiella trochoidea as high as in the red tide patch, suggesting that nutrients were not an initializing factor for this red tide. During the incubation of the red tide water sample without any nutrient addition, the phytoplankton biomass decreased gradually over 9 days. However, with a N addition, the phytoplankton biomass increased steadily until day 7, which suggested that nitrogen addition was able to sustain the high biomass of the red tide for a week with and without nutrients. In contrast, the red tide in the bay disappeared on the sampling day when the wind direction changed. These results indicated that initiation, maintenance and disappearance of the dinoflagellate Scrippsiella trochoidea red tide in the bay were not directly driven by changes in nutrients. Therefore, how nutrients are linked to the formation of red tides in coastal waters need to be further examined, particularly in relation to dissolved organic nutrients.     

Effect of nitrogen forms on growth,cell composition and N<Subscript>2</Subscript> fixation of <Emphasis Type="Italic">Cylindrospermopsis raciborskii</Emphasis> in phosphorus-limited chemostat cultures

The aim of this research was to test whether NH₄ ⁺ and NO₃ ⁻ affect the growth, P demand, cell composition and N₂ fixation of Cylindrospermopsis raciborskii under P limitation. Experiments were carried out in P-limited (200 μg l⁻¹ PO₄-P) chemostat cultures of C. raciborskii using an inflowing medium containing either 4,000 μg l⁻¹ NH₄-N, 4,000 μg l⁻¹ NO₃-N or no combined N. The results showed the cellular N:P and C:P ratios of C. raciborskii decreased towards the Redfield ratio with increasing dilution rate (D) due to the alleviation of P limitation. The cellular C:N and carotenoids:chlorophyll-a ratios also decreased with D, predominantly as a result of an increase in the chlorophyll-a and N content. The NH₄ ⁺ and NO₃ ⁻ supply reduced the P maintenance cell quota of C. raciborskii. Consequently, the biomass yield of the N₂-grown culture was significantly lower. The maximum specific growth rate of N₂-grown culture was also the lowest observed. It is suggested that these differences in growth parameters were caused by the P and energy requirement for heterocyte formation, nitrogenase synthesis and N₂ fixation. N₂ fixation was partially inhibited by NO₃ ⁻ and completely inhibited by NH₄ ⁺. It was probably repressed through the high N content of cells at high dissolved N concentrations. These results indicate that C. raciborskii is able to grow faster and maintain a higher biomass under P limitation where a sufficient supply of NH₄ ⁺ or NO₃ ⁻ is maintained. Information gained about the species-specific nutrient and pigment stoichiometry of C. raciborskii could help to access the degree of nutrient limitation in water bodies. Handling editor: Luigi Naselli-Flores     

Spatial and Temporal Distribution of Dissolved Inorganic Nutrients and Phytoplankton in Mida Creek,Kenya

The spatial and temporal distributions of dissolved inorganic nutrients were investigated between May 1996 and April 1997 in Mida Creek, a mangrove area along the north coast region of Kenya. The nutrient levels of pore water from boreholes/wells within the surrounding area of the creek were also investigated for comparison. In addition, phytoplankton distribution in Mida Creek was assessed in three stations within the creek in order to determine the structure and succession stages of the phytoplankton community and to provide an indication of the status of primary productivity of the creek. Measurements carried out within the creek revealed that the mean concentration ranges for NH₄⁺ –N, (NO ₂⁻ + NO₃⁻)−N, PO₄³⁻ −P and SiO₃²⁻ −Si were: 0.002–5.45; 0.12–5.63; 0.10–0.58 and 1.31–81.36 μM, respectively. For the case of boreholes/wells found in the surrounding area, their respective nutrient levels were found to lie in the ranges 0.4–907.0; 16.7–4897.0; 1.09–22.39 and 83.9–596.0 μM. A total of 295 species of phytoplankton belonging to 78 genera were identified with great temporal variability in abundance in all the stations sampled. The most dominant algal members in the Creek included Chaetoceros spp., Chroococcus limneticus and Oscillatoria spp. The diversity values recorded were indicative of mesotrophic conditions. The highest nutrient concentration levels within the creek were measured during the wet season as compared to dry season and this trend closely corresponded with that of the phytoplankton productivity. However, no significant variation ( p > 0.05) was found in all cases with respect to the tidal cycles. On the contrary, diurnal nutrient concentrations especially in areas with high flooding duration (>12 h) were found to be highest during the dry season as opposed to wet season for all nutrients except for SiO₃²⁻. The relatively high nutrient laden groundwater outflow into the creek water, coupled with surface runoff events during wet season, are the two main factors responsible for the elevated nutrients in the creek waters in the absence of river inflow into the creek.     

Ecological indicators of nutrient enrichment, freshwater wetlands, Midwestern United States (U.S.)

Vegetation and soil indicators of nutrient condition were evaluated in 30 wetlands, 10 each in 3 Nutrient Ecoregions (NE) (VI-Corn Belt and Northern Great Plains, VII-Mostly Glaciated Dairy Region, IX-Temperate Forested Plains and Hills) of the Midwestern United States (U.S.) to identify robust indicators for assessment of wetland nutrient enrichment and eutrophication. Nutrient condition was characterized by surface water inorganic N (NH₄-N, NO₃-N) and P (PO₄-P) concentrations measured seasonally for 1 year, plant available and total soil N and P, and aboveground biomass, leaf N and P and species composition of emergent vegetation measured at the end of the growing season. Aboveground biomass, nutrient uptake and species composition were positively related to surface water NH₄-N (N) but not to PO₄-P or NO₃-N. Aboveground biomass and biomass of aggressive species, Typha spp. plus Phalaris arundinacea, increased asymptotically with surface water N whereas leaf P, senesced leaf N and senesced leaf P increased linearly with N. And, species richness declined with surface water N. Soil total P was positively related to surface water PO₄-P but it was the only soil indicator related to wetland nutrient condition. Individual regressions for each NE generally were superior to a single regression for all NEs. In NE VI (Corn Belt), few indicators were related to surface water N because of the high degree of anthropogenic disturbance (85% of the landscape is cleared) as compared to NEs VII and IX (24–53% cleared). Of the indicators evaluated, stem height (r² = 0.42 for all NEs, r² = 0.56 for NE VII + IX) and percent biomass of aggressive species, Typha spp. plus Phalaris, (r² = 0.46 for all NEs, r² = 0.54 for NE VII + IX), were the best predictors of wetland nutrient enrichment. Vegetation-based indicators are a promising tool for assessment of wetland nutrient condition but they may not be effective in NEs where landscape disturbance is intense and widespread.     

Biofiltering efficiency, uptake and assimilation rates of Ulva clathrata (Roth) J. Agardh (Clorophyceae) cultivated in shrimp aquaculture waste water

The growth, biofiltering efficiency and uptake rates of Ulva clathrata were studied in a series of outdoor tanks, receiving waste water directly from a shrimp (Litopenaeus vannamei) aquaculture pond, under constant aeration and two different water regimes: (1) continuous flow, with 1 volume exchange a day (VE day^-1) and (2) static regime, with 1 VE after 4 days. Water temperature, salinity, pH, dissolved inorganic nitrogen (DIN), phosphate (PO₄), chlorophyll-a (chl-a), total suspended solids (TSS), macroalgal biomass (fresh weight) and tissue nutrient assimilation were monitored over 12 days. Ulva clathrata was highly efficient in removing the main inorganic nutrients from effluent water, stripping 70–82% of the total ammonium nitrogen (TAN) and 50% PO₄ within 15 h. Reductions in control tanks were much lower (Tukey HSD, P < 0.05). After 3 days, the mean uptake rates by the seaweed biomass under continuous flow were 3.09 mg DIN g DW day⁻¹ (383 mg DIN m⁻² day⁻¹) and 0.13 mg PO₄ g DW day⁻¹ (99 mg PO₄ m⁻² day⁻¹), being significantly higher than in the static regime (Tukey HSD, P < 0.05). The chl-a decreased in seaweed tanks, suggesting that U. clathrata inhibited phytoplankton growth. Correlations between the cumulative values of DIN removed from the water and total nitrogen assimilated into the seaweed biomass (r = 0.7 and 0.8, P < 0.05), suggest that nutrient removal by U. clathrata dominated over other processes such as phytoplankton and bacterial assimilation, ammonia volatilization and nutrient precipitation.     

Phytoplankton biomass and primary production in the marginal ice zone of the northwestern Weddell Sea during austral summer

During the austral summer of 1995, distributions of phytoplankton biomass (as chlorophyll a), primary production, and nutrient concentrations along two north-south transects in the marginal ice zone of the northwestern Weddell Sea were examined as part of the 8th Korean Antarctic Research Program. An extensive phytoplankton bloom, ranging from 1.6 to 11.2 mg m⁻³ in surface chlorophyll a concentration, was encountered along the eastern transect and extended ca. 180 km north of the ice edge. The spatial extent of the bloom was closely related to the density field induced by the input of meltwater from the retreating sea ice. However, the extent (ca. 200 km) of the phytoplankton bloom along the western transect exceeded the meltwater-influenced zone (ca. 18 km). The extensive bloom along the western transect was more closely related to local hydrography than to the proximity of the ice edge and the resulting meltwater-induced stability of the upper water column. In addition, the marginal ice zone on the western transect was characterized by a deep, high phytoplankton biomass (up to 8 mg Chl a m⁻³) extending to 100-m depth, and the decreased nutrient concentration, which was probably caused by passive sinking from the upper euphotic zone and in situ growth. Despite the low bloom intensity relative to the marginal ice zone in both of the transects, mean primary productivity (2.6 g C m⁻² day⁻¹) in shelf waters corresponding to the northern side of the western transect was as high as in the marginal ice zone (2.1 g C m⁻² day⁻¹), and was 4.8 times greater than that in open waters, suggesting that shelf waters are as highly productive as the marginal ice zone. A comparison between the historical productivity data and our data also shows that the most productive regions in the Southern Ocean are shelf waters and the marginal ice zone, with emerging evidence of frontal regions as another major productive site. Accepted: 27 September 1998     

An artificial neural network approach for studying phytoplankton succession

Artificial neural networks are used to model phytoplankton succession and gain insight into the relative strengths of bottom-up and top-down forces shaping seasonal patterns in phytoplankton biomass and community composition. Model comparisons indicate that patterns in chlorophyll aconcentrations response instantaneously to patterns in nutrient concentrations (phosphorous (P), nitrite and nitrate (NO₂/NO₃–N) and ammonium (NH₄–H) concentrations) and zooplankton biomass (daphnid cladocera and copepoda biomass); whereas lagged responses in an index of algal community composition are evident. A randomization approach to neural networks is employed to reveal individual and interacting contributions of nutrient concentrations and zooplankton biomass to predictions of phytoplankton biomass and community composition. The results show that patterns in chlorophyll aconcentrations are directly associated with P, NO₂/NO₃–N and daphnid cladocera biomass, as well as related to interactions between daphnid cladocera biomass, and NO₂/NO₃–N and P. Similarly, patterns in phytoplankton community composition are associated with NO₂/NO₃–N and daphnid cladocera biomass; however show contrasting patterns in nutrient– zooplankton and zooplankton–zooplankton interactions. Together, the results provide correlative evidence for the importance of nutrient limitation, zooplankton grazing and nutrient regeneration in shaping phytoplankton community dynamics. This study shows that artificial neural networks can provide a powerful tool for studying phytoplankton succession by aiding in the quantification and interpretation of the individual and interacting contributions of nutrient limitation and zooplankton herbivory on phytoplankton biomass and community composition under natural conditions.     

Nutrient limitation of phytoplankton communities in Subarctic lakes and ponds in Wapusk National Park, Canada

We determined the limiting nutrient of phytoplankton in 21 lakes and ponds in Wapusk National Park, Canada, using nutrient enrichment bioassays to assess the response of natural phytoplankton communities to nitrogen and phosphorus additions. The goal was to determine whether these Subarctic lakes and ponds were nutrient (N or P) limited, and to improve the ability to predict future impacts of increased nutrient loading associated with climate change. We found that 38% of lakes were not limited by nitrogen or phosphorus, 26% were co-limited by N and P, 26% were P-limited and 13% were N-limited. TN/TP, DIN/TP and NO₃ ⁻/TP ratios from each lake were compared to the Redfield ratio to predict the limiting nutrient; however, these predictors only agreed with 29% of the bioassay results, suggesting that nutrient ratios do not provide a true measure of nutrient limitation within this region. The N-limited lakes had significantly different phytoplankton community composition with more chrysophytes and Anabaena sp. compared to all other lakes. N and P limitation of phytoplankton communities within Wapusk National Park lakes and ponds suggests that increased phytoplankton biomass may result in response to increased nutrient loading associated with environmental change.     

The role of phosphorus on planktonic production of the Gironde plume waters in the Bay of Biscay

More and more studies emphasize the status of phosphorus (P)as the principal limiting nutrient of phytoplankton growth,especially in coastal waters under the influence of freshwaterdischarges. The purpose of the present paper is to investigatethe role of P on planktonic production in the waters influencedby the Gironde discharges; the Gironde being one of the twolargest rivers on the French Atlantic coast. The survey is basedon several cruises made in 1998 and 1999. Two different patternswere observed for waters with salinity below and above 34.5.For waters with salinity < 34.5, P was found to be the firstlimiting nutrient of winter and spring phytoplankton blooms,based on undetectable phosphate (< 20 nM), high NO₃ : PO₄ratios, typically > 100 : 1, short phosphate turnover time(1 to 2 h), high alkaline phosphatase activities (mean of 176nM h^-1 in late May 1999) and ultimately great increases of chlorophylla (Chl a) and primary production in phosphate-enriched samplesrelative to controls. This limitation could be partly explainedby the Gironde nutrient supplies, which were phosphate deficientcompared with the mineral nitrogen(N_min : PO₄ was > 40 withina salinity range 16–33). In summer, corresponding to theperiod of low influence of Gironde supplies (low runoff anda spreading effect of the plume), phytoplankton growth wouldbe controlled by both P and nitrogen (N), according to low nitrateand the major effect of combined P+N (NH₄) enrichment on Chla and primary production compared with the addition of N orP singly. In early October, after the first autumn gales, themixed layer was enriched with a sufficient supply of nutrientsto support exponential phytoplankton growth for 4 days in enclosures.The pattern was different for waters at the limit of the Girondeplume and Atlantic oceanic waters (within salinity range 34.5–35.4).P would not be the single limiting nutrient of winter bloomsand spring phytoplankton growth since low phosphate, and alsolow nitrate and silicate, availability were recorded and phosphateaddition alone had no effect on phytoplankton biomass and productionin bioassays. The early P limitation of winter blooms had consequencesfor the phytoplankton community structure in the Gironde plumewaters (salinity < 34.5). Whereas major cells of these bloomswere greater than 20 µm in size, phytoplankton growthin spring and autumn was dominated by 3–20 µm (e.g.53% of Chl a in late April 1999) and < 3 µm cells (e.g.29% of Chl a). The decreasing size of phytoplankton cells isemphasized by the severe competition between bacteria and algaefor phosphate, since bacteria dominated phosphate uptake inspring (e.g. 87% in late April, 77% in late May). Bacteria tendedto have greater affinity for phosphate and seemed also to beP limited at certain times in spring, according to results fromphosphate enrichment bioassays in late May 1999. The alternativemethod for phytoplankton to obtain P would be the use of thedissolved organic phosphorus pool by alkaline phosphatase activity.According to the movement of ³³P after initial labelling ofmicrobial populations and a subsequent cold chase, the majortransfer of P occurred from the bacterial to the dissolved fraction.We hypothesize that algae obtain part of its dissolved organicphosphorus from bacteria-originated organic phosphorus compounds.     

Seasonal variation in nutrient limitation of microbial biofilms colonizing organic and inorganic substrata in streams

Humans have increased the availability of nutrients including nitrogen and phosphorus worldwide; therefore, understanding how microbes process nutrients is critical for environmental conservation. We examined nutrient limitation of biofilms colonizing inorganic (fritted glass) and organic (cellulose sponge) substrata in spring, summer, and autumn in three streams in Michigan, USA. Biofilms were enriched with nitrate (NO₃ ⁻), phosphate (PO₄ ³⁻), ammonium (NH₄ ⁺), NO₃ ⁻ + PO₄ ³⁻, NH₄ ⁺ + PO₄ ³⁻, or none (control). We quantified biofilm structure and function as chlorophyll a (i.e., primary producer biomass) and community respiration on all substrata. In one stream, we characterized bacterial and fungal communities on cellulose in autumn using clone library sequencing and denaturing gradient gel electrophoresis to determine if community structure was linked to nutrient limitation status. Despite oligotrophic conditions, primary producer biomass was infrequently nutrient limited. In contrast, respiration on organic substrata was frequently limited by N + P combinations. We found no difference between biofilm response to NH₄ ⁺ versus NO₃ ⁻ enrichment, although the response to both N-species was positively related to water column PO₄ ³⁻ concentrations and temperature. Molecular analysis for fungal community composition suggested no relationship to nutrient limitation, but the dominant members of the bacterial community on cellulose were different on NO₃ ⁻, PO₄³, and NO₃ ⁻ + PO₄ ³⁻ treatments relative to control, NH₄ ⁺, and NH₄ ⁺ + PO₄ ³⁻ treatments, which matched patterns for biofilm respiration rates from each treatment. Our results show discrete patterns of nutrient limitation dependent upon substratum type and season, and imply changes in bacterial community structure and function may be linked following nutrient enrichment in streams.     

Periphyton nitrogenase activity as an indicator of wetland eutrophication: spatial patterns and response to phosphorus dosing in a northern Everglades ecosystem

The use of periphyton nitrogenase activity (biological N₂ fixation) as an indicator of wetland P impact was assessed using patterns of nutrient content (C, N, P, Ca, Mg, K, Fe, and Mn) and acetylene reduction (AR) in floating cyanobacterial periphyton mat (metaphyton) communities of a P-enriched portion of the Florida Everglades, USA (Water Conservation Area-2A, WCA-2A). Spatial patterns of nutrients indicate the enrichment of floating mat periphyton N, P, Fe, and K, and the reduction of Mn and TN:TP in enriched marsh areas. In highly enriched areas, floating mat periphyton AR was approximately threefold greater than that in less enriched, interior marsh zones. Multiple regression models indicated AR dependence on P in eutrophic WCA-2A areas while the AR of more interior marsh periphyton mats was more closely related to tissue levels of Ca and Fe. Nitrogenase activity of floating mat periphyton from P-loaded mesocosms revealed a significant enhancement of N₂ fixation in samples receiving approximately 2–3 mg P m⁻² of cumulative P dosing or with biomass TP content of 100–300 mg kg⁻¹. At P contents above the optimum, mat periphyton AR was suppressed possibly as a result of changes in species composition or increased levels of NH₄⁺. After 3 years of dosing, consistently high AR occurred only at low rates of P enrichment (0.4–0.8 g P m⁻² yr⁻¹), and the patterns appeared to be seasonal. These findings agree with the hypothesis that P availability is a key determinant of nitrogenase activity in aquatic systems, and thus, may support the use of periphyton nitrogenase to indicate P impacts in P-limited systems. These results also demonstrate the potential existence of a P threshhold for biogeochemical alteration of periphyton mat function in the Everglades, and that cumulative loading of limiting nutrients (i.e., P), rather than instantaneous concentrations, should be considered when evaluating nutrient criteria.     

A mass balance evaluation of the ecological significance of historical nitrogen fluxes in Lake Kinneret

Lake Kinneret (LK) is a monomictic lake that has undergone significant biological and chemical changes over the last three decades of the twentieth century. The transition between the 1970s and the 1980s attracted a lot of scientific attention as it was marked by significant changes in the ecology of the lake. In the early 1980s, phytoplankton biomass increased, apparently in response to an increase in the external soluble reactive phosphorus (SRP) load. This period was marked by a rise in hypolimnetic levels of ammonium (NH₄) and SRP as well as surface water dissolved oxygen (DO) and pH. Cconcomitantly, in surface waters in winter levels of NH₄ increased and NO₃ decreased. In this study interrelationships amongst these observations were examined with a mass balance modelling approach, including simulation of individual nutrient sources and sinks, focusing on nitrogen fluxes in winter. The step-like rise in phytoplankton biomass in 1981 may have been triggered by the increase in winter external loads of SRP, as P is likely to be the growth-limiting nutrient during this season. The additional P load led to a sequence of changes including greater summer phytoplankton biomass, followed by enhanced sedimentation of organic matter. Furthermore, higher organic matter mineralization fluxes within the hypolimnion resulted in elevated levels of NH₄ and SRP in this layer through the 1980s, with a feedback to productivity in the trophogenic zone following seasonal destratification in early winter. In an apparent transition period (late 1970s to early 1980s), an increase in the modelled rate of nitrate (NO₃) production occurred via nitrification together with increased uptake of the additional nitrate by phytoplankton. These results are consistent with increased phytoplankton abundance and elevated levels of surface water NH₄ and DO during this period. Through this period the increase in phytoplankton uptake of NO₃ predominated over the increase in nitrification, and NO₃ concentrations in the 1980s were reduced compared with the previous decade, with increased partitioning of N in biomass and NH₄.     

Distribution of nitrogen-15 tracers applied to the canopy of a mature spruce-hemlock stand,Howland, Maine,USA

In N-limited ecosystems, fertilization by N deposition may enhance plant growth and thus impact C sequestration. In many N deposition–C sequestration experiments, N is added directly to the soil, bypassing canopy processes and potentially favoring N immobilization by the soil. To understand the impact of enhanced N deposition on a low fertility unmanaged forest and better emulate natural N deposition processes, we added 18 kg N ha⁻¹ year⁻¹ as dissolved NH₄NO₃ directly to the canopy of 21 ha of spruce-hemlock forest. In two 0.3-ha subplots, the added N was isotopically labeled as ¹⁵NH₄ ⁺ or ¹⁵NO₃ ⁻ (1% final enrichment). Among ecosystem pools, we recovered 38 and 67% of the ¹⁵N added as ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻, respectively. Of ¹⁵N recoverable in plant biomass, only 3–6% was recovered in live foliage and bole wood. Tree twigs, branches, and bark constituted the most important plant sinks for both NO₃ ⁻ and NH₄ ⁺, together accounting for 25–50% of ¹⁵N recovery for these ions, respectively. Forest floor and soil ¹⁵N retention was small compared to previous studies; the litter layer and well-humified O horizon were important sinks for NH₄ ⁺ (9%) and NO₃ ⁻ (7%). Retention by canopy elements (surfaces of branches and boles) provided a substantial sink for N that may have been through physico-chemical processes rather than by N assimilation as indicated by poor recoveries in wood tissues. Canopy retention of precipitation-borne N added in this particular manner may thus not become plant-available N for several years. Despite a large canopy N retention potential in this forest, C sequestration into new wood growth as a result of the N addition was only ~16 g C m⁻² year⁻¹ or about 10% above the current net annual C sequestration for this site.     

The effects of human activities and extreme meteorological events on sediment nitrogen dynamics in an urban estuary,Escambia Bay,Florida, USA

This study examined how sediment-sorbed PCBs and several large storms affected sediment nutrient dynamics based on potential nitrification rates and benthic flux measurements. PCBs were hypothesized to negatively affect potential nitrification rates due to the sensitivity of nitrifying bacteria. Sediment disturbance caused by the succession of storms, which can enhance nutrient inputs and phytoplankton production, was hypothesized to enhance both potential nitrification rates and benthic flux measurements as a result of higher nutrient and organic matter concentrations. Potential nitrification rates, benthic fluxes (NO₃ ⁻ + NO₂ ⁻, NH₄ ⁺, and DIP), sediment PCB content, water content, organic content, salinity, bottom water dissolved oxygen, and sediment chlorophyll were measured at 13 different sites in Escambia Bay during the summer of 2005. Potential nitrification rates were highest at deep, organic-rich sites. Total PCB content did not have a direct effect on potential nitrification rates. An analysis of recent changes in benthic processes in relation to extreme meteorological events was performed by comparing the 2005 results with data from 2000, 2003, and 2004. Storm effects on sediment biogeochemistry were mixed with sediment nitrogen dynamics enhanced at some sites but not others. In addition, SOC and NH₄ ⁺ fluxes increased in deeper channel sites after Hurricanes Ivan and Dennis, which could be attributed to the deposition of phytoplankton blooms. Sediment nutrient dynamics in Escambia Bay appear to be resilient to these extreme meteorological events since there were no significant effects on sediment processes in the Bay as a whole. Handling editor: P. Viaroli     

Phytoplankton production after the collapse of the Larsen A Ice Shelf,Antarctica

Part of the Larsen A Ice Shelf (64°15′S to 74°15′S) collapsed during January 1995. A first oceanographic and biological data set from the newly free waters was obtained during December 1996. Typical shelf waters with temperatures near and below the freezing point were found. A nutrient-rich water mass (max: PO₄ ³⁻ 1.80 μmol L⁻¹ and NO₃ ⁻ 27.64 μmol L⁻¹) was found between 70 and 200 m depth. Chlorophyll-a (Chl-a) values (max 14.24 μg L⁻¹) were high; surface oxygen saturation ranged between 86 and 148%. Diatoms of the genera Nitzschia and Navicula and the prymnesiophyte Phaeocystis sp. were the most abundant taxa found. Mean daily primary production (Pc) estimated from nutrient consumption was 14.80 ± 0.17 mgC m⁻³ day⁻¹. Pc was significantly correlated with total diatom abundance and Chl-a. Calculated ΔpCO₂ (difference of the CO₂ partial pressure between surface seawater and the atmosphere) was –30.5 μatm, which could have contributed to a net CO₂ flux from the atmosphere to the sea and suggests the area has been a CO₂ sink during the studied period. High phytoplankton biomass and production values were found in this freshly open area, suggesting its importance for biological CO₂ pumping.     

Benthic nutrient fluxes in a eutrophic,polymictic lake

Sediment release rates of soluble reactive phosphorus (SRP) and ammonium (NH₄) were determined seasonally at three sites (water depth 7, 14 and 20 m) in Lake Rotorua using in situ benthic chamber incubations. Rates of release of SRP ranged from 2.2 to 85.6 mg P m⁻² d⁻¹ and were largely independent of dissolved oxygen (DO) concentration. Two phases of NH₄ release were observed in the chamber incubations; high initial rates of up to 2,200 mg N m⁻² d⁻¹ in the first 12 h of deployment followed by lower rates of up to 270 mg N m⁻² d⁻¹ in the remaining 36 h of deployment. Releases of SRP and NH₄ were highest in summer and at the deepest of the three sites. High organic matter supply rates to the sediments may be important for sustaining high rates of sediment nutrient release. A nutrient budget of Lake Rotorua indicates that internal nutrient sources derived from benthic fluxes are more important than external nutrient sources to the lake.     

Microphytobenthos production in the Gulf of Fos, French Mediterranean coast

Microphytobenthic oxygen production was studied in the Gulf of Fos (French Mediterranean coast) during 1991/1992 using transparent and dark benthic chambers. Nine stations were chosen in depths ranging from 0.5 to 13 m, which represents more than 60% of bottoms in the Gulf. Positive net microphytobenthic oxygen production was seasonally detected down to 13 m; the maximum value attained was 60 mg O₂ m⁻² h⁻¹ (0.7–0.8 g O₂ m⁻² d⁻¹) in sediments at 0.5 m depth during spring and winter. Respiration rates were maximum in the sediments located at the mussel farm (5 m), in the center of the Gulf, with 135 mg O₂ m⁻² h⁻¹ in spring (3.2 g O₂ m⁻² d⁻¹); in the other locations, it ranged from 3.3 to 58.2 mg O₂ m⁻² h⁻¹ (0.08–1.4 g O₂ m⁻² d⁻¹). Compared to phytoplankton, microphytobenthos production was higher only in the bottoms < 1 m depth. In deeper bottom waters, phytoplankton production could be absent due to light limitation, while microphytobenthos was still productive. Phytoplankton production m⁻² was generally higher than microphytobenthic production. Microphytobenthic biomass, higher than phytoplanktonic, varied from 27 to 379 mg Chl a m⁻², the maximum in the mussel farm sediments, with the minimum in sandy shallow bottoms. Pigment analysis showed that microphytobenthos consisted mainly of diatoms (Chl c and fucoxanthin) but other algal groups containing Chl b could become seasonally important. A Principal Component Analysis suggested that the main statistical factors explaining the distribution of our observations may be interpreted in terms of enrichment in phaeopigments and light; the role of Chl a appearing paradoxically as secondary in benthic production rates. Phaeopigments are mainly constituted by phaeophorbides, which indicate grazing processes. The influence of the mussel farm on the oxygen balance is noticeable in the whole Gulf.