Wayne S. Gardner

Controlling harmful cyanobacterial blooms in a hyper-eutrophic lake (Lake Taihu, China): the need for a dual nutrient (N & P) management strategy.

Harmful cyanobacterial blooms, reflecting advanced eutrophication, are spreading globally and threaten the sustainability of freshwater ecosystems. Increasingly, non-nitrogen (N(2))-fixing cyanobacteria (e.g., Microcystis) dominate such blooms, indicating that both excessive nitrogen (N) and phosphorus (P) loads may be responsible for their proliferation. Traditionally, watershed nutrient management efforts to control these blooms have focused on reducing P inputs. However, N loading has increased dramatically in many watersheds, promoting blooms of non-N(2) fixers, and altering lake nutrient budgets and cycling characteristics. We examined this proliferating water quality problem in Lake Taihu, Chinas 3rd largest freshwater lake. This shallow, hyper-eutrophic lake has changed from bloom-free to bloom-plagued conditions over the past 3 decades. Toxic Microcystis spp. blooms threaten the use of the lake for drinking water, fisheries and recreational purposes. Nutrient addition bioassays indicated that the lake shifts from P limitation in winter-spring to N limitation in cyanobacteria-dominated summer and fall months. Combined N and P additions led to maximum stimulation of growth. Despite summer N limitation and P availability, non-N(2) fixing blooms prevailed. Nitrogen cycling studies, combined with N input estimates, indicate that Microcystis thrives on both newly supplied and previously-loaded N sources to maintain its dominance. Denitrification did not relieve the lake of excessive N inputs. Results point to the need to reduce both N and P inputs for long-term eutrophication and cyanobacterial bloom control in this hyper-eutrophic system.

Sediment Trap Studies in Lake Michigan: Resuspension and Chemical Fluxes in the Southern Basin

The results of 4 years (1977–80) of sediment trap sample collection in the southeastern region of Lake Michigan are summarized and compared with water column and sediment characteristics. Mass flux data indicate strong seasonal patterns, with maximum fluxes recorded during the unstratified period. The large amount of winter resuspension is a mechanism which provides an intimate coupling of recent sediments and the water column. Also these trap study results indicate that there is a near-bottom (10-m-thick) benthic nepheloid layer whose chemical composition approaches that of deep water (fine grain) sediments. The amount of resuspended NaOH extractable phosphorus injected into the euphotic zone is estimated as approximately equal to the load of new phosphorus entering southern Lake Michigan. The role of resuspension in the cycling of organic carbon and contaminants associated with it appears to be important.

Ecosystem-Level Effects of Zebra Mussels (Dreissena polymorpha): An Enclosure Experiment in Saginaw Bay, Lake Huron

We examined the short-term effects of zebra mussels (Dreissena polymorpha) on ecosystem processes in late August 1991 in Saginaw Bay, Lake Huron. Four 1,600-L enclosures, made of Fabreen with a diameter of 1 m, a depth of 2 m, and closed at the bottom, were used to enclose natural plankton communities. These communities were dominated by diatoms with some chlorophytes, chrysophytes, and cyanophytes. Phytoplankton growth was limited by P-availability. Two enclosures were held as controls, and zebra mussels encrusting unionid shells were suspended in two of the enclosures: one enclosure (HZ) contained approximately four-fold greater numbers of mussels than the other (LZ). The concentration of suspended particles, chlorophyll, and algal biomass in HZ and LZ declined over a 6-day interval. Diatom numbers declined more than other taxa. Phytoplankton growth rates in HZ and LZ increased to near μmax; there was no apparent change in photosynthetic parameters a or Pmax scaled for chlorophyll. Soluble reactive P (SRP) increased significantly (p < 0.05) in HZ but not LZ. Dissolved organic P (DOP) and ammonium ion were elevated; dissolved organic carbon (DOC) was unchanged in HZ and LZ. The rate of phosphate uptake by bacteria and algae declined to less than 2% of controls; this rate decrease could not be explained simply by grazing losses or isotope dilution. The rate of ammonium regeneration by the plankton and the potential rate of ammonium uptake by the plankton did not differ significantly in HZ or LZfrom the control enclosures. Our findings indicate that the zebra mussel can have significant short-term effects on phytoplankton abundance, water transparency, water chemistry and phosphorus dynamics. We propose a model of zebra mussel effects that suggests high densities of zebra mussels may indirectly alter and control those processes that are rate-limited or concentration-limited by nutrient availability.

The effect of salinity on ammonium sorption in aquatic sediments: Implications for benthic nutrient recycling

Ambient exchangeable ammonium concentrations in freshwater sediments are generally considerably greater than those reported for marine sediments. Laboratory measurements indicate that competition for cation exchange sites by ions in seawater is a factor responsible for the lower exchangeable ammonium concentrations in marine sediments. Exchangeable ammonium concentrations were 3- to 6-fold higher when river and estuarine sediments were incubated with fresh water relative to the same sediments incubated with salt water (%.-23). A model was developed to explore the implications for benthic nitrogen cycling of this salinity effect on exchangeable ammonium concentrations. Ammonium diffusion, exchangeable and dissolved ammonium concentrations, and nitrification rates were components of the model formulation. The model output suggests that higher exchangeable ammonium concentrations predicted in fresh water relative to marine sediments can markedly increase the fraction of the ammonium produced in sediments that is nitrified (and subsequently denitrified). These results are consistent with field and experimental laboratory data which indicate that a larger percentage of net ammonium production in aerobic freshwater sediments is nitrified and denitrified (80–100%) relative to marine sediments (40–60%).

Simultaneous Measurement of Denitrification and Nitrogen Fixation Using Isotope Pairing with Membrane Inlet Mass Spectrometry Analysis

ABSTRACT A method for estimating denitrification and nitrogen fixation simultaneously in coastal sediments was developed. An isotope-pairing technique was applied to dissolved gas measurements with a membrane inlet mass spectrometer (MIMS). The relative fluxes of three N2 gas species (28N2,29N2, and 30N2) were monitored during incubation experiments after the addition of15NO3−. Formulas were developed to estimate the production (denitrification) and consumption (N2 fixation) of N2 gas from the fluxes of the different isotopic forms of N2. Proportions of the three isotopic forms produced from15NO3− and14NO3− agreed with expectations in a sediment slurry incubation experiment designed to optimize conditions for denitrification. Nitrogen fixation rates from an algal mat measured with intact sediment cores ranged from 32 to 390 μg-atoms of N m−2 h−1. They were enhanced by light and organic matter enrichment. In this environment of high nitrogen fixation, low N2 production rates due to denitrification could be separated from high N2 consumption rates due to nitrogen fixation. Denitrification and nitrogen fixation rates were estimated in April 2000 on sediments from a Texas sea grass bed (Laguna Madre). Denitrification rates (average, 20 μg-atoms of N m−2 h−1) were lower than nitrogen fixation rates (average, 60 μg-atoms of N m−2 h−1). The developed method benefits from simple and accurate dissolved-gas measurement by the MIMS system. By adding the N2 isotope capability, it was possible to do isotope-pairing experiments with the MIMS system.

Nitrogen dynamics and microbial food web structure during a summer cyanobacterial bloom in a subtropical, shallow, well-mixed, eutrophic lake (Lake Taihu, China)

Nitrogen dynamics and microbial food web structure were characterized in subtropical, eutrophic, large (2,338 km2), shallow (1.9 m mean depth), and polymictic Lake Taihu (China) in Sept–Oct 2002 during a cyanobacterial bloom. Population growth and industrialization are factors in trophic status deterioration in Lake Taihu. Sites for investigation were selected along a transect from the Liangxihe River discharge into Meiliang Bay to the main lake. Water column nitrogen and microbial food web measurements were combined with sediment-water interface incubations to characterize and identify important processes related to system nitrogen dynamics. Results indicate a gradient from strong phosphorus limitation at the river discharge to nitrogen limitation or co-limitation in the main lake. Denitrification in Meiliang Bay may drive main lake nitrogen limitation by removing excess nitrogen before physical transport to the main lake. Five times higher nutrient mineralization rates in the water column versus sediments indicate that sediment nutrient transformations were not as important as water column processes for fueling primary production. However, sediments provide a site for denitrification, which, along with nitrogen fixation and other processes, can determine available nutrient ratios. Dissimilatory nitrate reduction to ammonium (DNRA) was important, relative to denitrification, only at the river discharge site, and nitrogen fixation was observed only in the main lake. Reflecting nitrogen cycling patterns, microbial food web structure shifted from autotrophic (phytoplankton dominated) at the river discharge to heterotrophic (bacteria dominated) in and near the main lake.

Effects of the Zebra Mussel (Dreissena polymorpha Pallas) on Protozoa and Phytoplankton from Saginaw Bay, Lake Huron

Direct effects of the grazing activities of the zebra mussel, Dreissena polymorpha, on the natural assemblage of planktonic protozoa and algae from Saginaw Bay, Lake Huron, were studied in September and October 1994. Water and mussels collected from two eutrophic sites were incubated in an outdoor “natural light” incubator at ambient temperature for 24 hours. Experiments were conducted in 4-L bottles with screened (40 or 53-μm net) or unscreened water and with and without mussels. Despite relatively high growth rates of protozoa on both dates, mussels lowered protozoan numbers by 70–80% and reduced the species richness of the protozoan community by 30–50%. Large heterotrophic flagellates were reduced up to 100% while peritrichous ciliates attached to the colonies of blue-greens were reduced only by 50%. Dreissena selectively removed nanoplanktonic Cryptomonas and Cyclotella, but had no significant effect on the predominant phytoplankton species, Microcystis. Overall, Dreissena clearance rates were low in the presence of this cyanophyte species. We conclude that zebra mussels, in regions where they are abundant, can cause significant changes in composition of both the protozoan and phytoplankton communities.

The effects of sea salts on the forms of nitrogen released from estuarine and freshwater sediments: Does ion pairing affect ammonium flux?

In sediments with oxidized surface layers, the percentage of mineralized nitrogen that is nitrified/denitrified, compared with that released directly as ammonium, appears to be affected by the presence of sea salts. In estuarine systems, a significant portion of the nitrogen is released as ammonium, whereas in freshwater systems, most of the mineralized nitrogen is often released from the sediments as nitrogen gas. We hypothesized that this discrepancy is caused by differential competition between physical diffusion and nitrification/denitrification in the two systems. The vertical migration (by Fickian diffusion) of ammonium out of the oxic layer may be hindered by cation exchange (or sorption) interactions with sediment particles to a greater extent in fresh water than in estuarine systems. The resulting relatively long residence time, and potentially high levels of particle-bound ammonium in the freshwater sediments, would favor nitrification as the major ammonium removal process. By contrast, ion pair formation of ammonium with seawater anions and blockage of sediment cation exchange sites with seawater cations may allow a sizable fraction of the ammonium to diffuse out of estuarine sediments before it is nitrified. A salt effect, consistent with this hypothesis, has been demonstrated in experimental systems by changing the ionic composition of water flowing above intact cores of freshwater and estuarine sediments. Steady-state ammonium release from Lake Michigan sediments was substantially enhanced in the presence of 30% seawater over that in the presence of lake water alone. Likewise, steady-state ammonium release, from Ochlockonee River and Bay sediments (Florida) and from Toms River and Barnegat Bay sediments (New Jersey), was usually higher in the presence of diluted synthetic seawater than it was in the presence of fresh water.

A model study of the coupled biological and physical dynamics in Lake Michigan

A coupled physical and biological model was developed for Lake Michigan. The physical model was the Princeton ocean model (POM) driven directly by observed winds and net surface heat flux. The biological model was an eight-component, phosphorus-limited, lower trophic level food web model, which included phosphate and silicate for nutrients, diatoms and non-diatoms for dominant phytoplankton species, copepods and protozoa for dominant zooplankton species, bacteria and detritus. Driven by observed meteorological forcings, a 1-D modeling experiment showed a controlling of physical processes on the seasonal variation of biological variables in Lake Michigan: diatoms grew significantly in the subsurface region in early summer as stratification developed and then decayed rapidly in the surface mixed layer when silicate supplied from the deep stratified region was reduced as a result of the formation of the thermocline. The non-diatoms subsequently grew in mid and late summer under a limited-phosphate environment and then declined in the fall and winter as a result of the nutrient consumption in the upper eutrophic layer, limitation of nutrients supplied from the deep region and meteorological cooling and wind mixing. The flux estimates suggested that the microbial loop had a significant contribution in the growth of microzooplankton and hence, to the lower-trophic level food web system. The model results agreed with observations, suggesting that the

A Review of Water Column Processes Influencing Hypoxia in the Northern Gulf of Mexico

In this review, we use data from field measurements of biogeochemical processes and cycles in the Mississippi River plume and in other shelf regions of the northern Gulf of Mexico to determine plume contributions to coastal hypoxia. We briefly review pertinent findings from these process studies, review recent mechanistic models that synthesize these processes to address hypoxia-related issues, and reinterpret current understanding in the context of these mechanistic models. Some of our conclusions are that both nitrogen and phosphorus are sometimes limiting to phytoplankton growth; respiration is the main fate of fixed carbon in the plume, implying that recycling is the main fate of nitrogen; decreasing the river nitrate loading results in less than a 1:1 decrease in organic matter sinking from the plume; and sedimenting organic matter from the Mississippi River plume can only fuel about 23% of observed coastal hypoxia, suggesting significant contributions from the Atchafalaya River and, possibly, coastal wetlands. We also identify gaps in our knowledge about controls on hypoxia, and indicate that some reinterpretation of our basic assumptions about this system is required. There are clear needs for improved information on the sources, rates, and locations of organic matter sedimentation; for further investigation of internal biogeochemical processes and cycling; for improved understanding of the rates of oxygen diffusion across the pycnocline; for identification and quantification of other sources of organic matter fueling hypoxia or other mechanisms by which Mississippi River derived organic matter fuels hypoxia; and for the development of a fully coupled physical-biogeochemical model.