Hypoxia in the northern Gulf of Mexico: Does the science support the Plan to Reduce,Mitigate, and Control Hypoxia?

We update and reevaluate the scientific information on the distribution, history, and causes of continental shelf hypoxia that supports the 2001 Action Plan for Reducing, Mitigating, and Controlling Hypoxia in the Northern Gulf of Mexico (Mississippi River/Gulf of Mexico Watershed Nutrient Task Force 2001), incorporating data, publications, and research results produced since the 1999 integrated assessment. The metric of mid-summer hypoxic area on the LouisianaTexas shelf is an adequate and suitable measure for continued efforts to reduce nutrients loads from the Mississippi River and hypoxia in the northern Gulf of Mexico as outlined in the Action Plan. More frequent measurements of simple metrics (e.g., area and volume) from late spring through late summer would ensure that the metric is representative of the system in any given year and useful in a public discourse of conditions and causes. The long-term data on hypoxia, sources of nutrients, associated biological parameters, and paleoindicators continue to verify and strengthen the relationship between the nitratenitrogen load of the Mississippi River, the extent of hypoxia, and changes in the coastal ecosystem (eutrophication and worsening hypoxia). Multiple lines of evidence, some of them representing independent data sources, are consistent with the big picture pattern of increased eutrophication as a result of long-term nutrient increases that result in excess carbon production and accumulation and, ultimately, bottom water hypoxia. The additional findings arising since 1999 strengthen the science supporting the Action Plan that focuses on reducing nutrient loads, primarily nitrogen, through multiple actions to reduce the size of the hypoxic zone in the northern Gulf of Mexico.     

Control of the variability of nutrients and suspended sediments in a gulf coast estuary by climatic forcing and spring discharge of the Atchafalaya River

Water column nutrients (nitrate, ammonium, soluble reactive phosphate, total Kjeldahl nitrogen, and total phosphorus) and suspended sediments (SS) were measured during one 44-h and two 28-h periods in March 1982 at two stations in Fourleague Bay, Louisiana, which is located at the mouth of the Atchafalaya River, a distributary of the Mississippi River. River water (a source of nitrate, total Kjeldahl nitrogen, total phosphorus, and suspended sediments to Fourleague Bay) flows into the upper reaches of the bay during high tide and frontal overrunning conditions with northerly and westerly winds. During one sampling period, decreasing wind speed and the rising tide resulted in Atchafalaya River water inundating the bay and nitrate concentrations in the upper bay increasing from 30–70 μM to 90–118 μM. Significant variations in nutrients associated with the movement of water masses from the river, marshes, and Gulf of Mexico occurred over several different time scales. Tidal transport occurred over 25-h periods, while frontal passages occurred at 3-d to 5-d intervals. Variability in nutrient and suspended sediment concentrations over these relatively short time scales can be as great as seasonal variability in the bay.     

A preliminary mass balance model of primary productivity and dissolved oxygen in the Mississippi River Plume/Inner Gulf Shelf Region

A deterministic, mass balance model for phytoplankton, nutrients, and dissolved oxygen was applied to the Mississippi River Plume/Inner Gulf Shelf (MRP/IGS) region. The model was calibrated to a comprehensive set of field data collected during July 1990 at over 200 sampling stations in the northern Gulf of Mexico. The spatial domain of the model is represented by a three-dimensional, 21-segment water-column grid extending from the Mississippi River Delta west to the Louisiana-Texas border, and from the shoreline seaward to the 30–60 m bathymetric contours. Diagnostic analyses and numerical experiments were conducted with the calibrated model to better understand the environmental processes controlling primary productivity and dissolved oxygen dynamics in the MRP/IGS region. Underwater light attenuation appears relatively more important than nutrient limitation in controlling rates of primary productivity. Chemical-biological processes appear relatively more important than advective-dispersive transport processes in controlling bottom-water dissolved oxygen dynamics. Oxidation of carbonaceous material in the water column, phytoplankton respiration, and sediment oxygen demand all appear to contribute significantly to total oxygen depletion rates in bottom waters. The estimated contribution of sediment oxygen demand to total oxygen-depletion rates in bottom waters ranges from 22% to 30%. Primary productivity appears to be an important source of dissolved oxygen to bottom waters in the region of the Atchafalaya River discharge and further west along the Louisiana Inner Shelf. Dissolved oxygen concentrations appear very sensitive to changes in underwater light attenuation due to strong coupling between dissolved oxygen and primary productivity in bottom waters. The Louisiana Inner Shelf in the area of the Atchafalaya River discharge and further west to the Texas border appears to be characterized by significantly different light attenuation-depth-primary productivity relationships than the area immediately west of the Mississippi Delta. Nutrient remineralization in the water column appears to contribute significantly to maintaining chlorophyll concentrations on the Louisiana Inner Shelf.     

Nutrient changes in the Mississippi River and system responses on the adjacent continental shelf

The Mississippi River system ranks among the world's top 10 rivers in freshwater and sediment inputs to the coastal ocean. The river contributes 90% of the freshwater loading to the Gulf of Mexico, and terminates amidst one of the United States' most productive fisheries regions and the location of the largest zone of hypoxia, in the western Atlantic Ocean. Significant increases in riverine nutrient concentrations and loadings of nitrate and phosphorus and decreases in silicate have occurred this century, and have accelerated since 1950. Consequently, major alterations have occurred in the probable nutrient limitation and overall stoichiometric nutrient balance in the adjacent continental shelf system. Changes in the nutrient balances and reduction in riverine silica loading to, the continental shelf appear to have led to phytoplankton species shifts offshore and to an increase in primary production. The phytoplankton community response, as indicated by long-term changes in biological uptake of silicate and accumulation of biologically bound silica in sediments, has shown how the system has responded to changes in riverine nutrient loadings. Indeed, the accumulation of biologically bound silica in sediments beneath the Mississippi River plume increased during the past two decades, presumably in response to, increased nitrogen loading. The duration, size, and severity of hypoxia has probably increased as a consequence of the increased primary production. Management alternatives directed at water pollution issues within the Mississippi River watershed may have unintended and contrasting impacts on the coastal waters of the northern Gulf of Mexico.     

Going West: Nutrient Limitation of Primary Production in the Northern Gulf of Mexico and the Importance of the Atchafalaya River

To investigate controls on phytoplankton production along the Louisiana coastal shelf, we mapped salinity, nutrient concentrations (dissolved inorganic nitrogen (DIN) and phosphorus (P_i), silicate (Si)), nutrient ratios (DIN/P_i), alkaline phosphatase activity, chlorophyll and ¹⁴C primary productivity on fine spatial scales during cruises in March, May, and July 2004. Additionally, resource limitation assays were undertaken in a range of salinity and nutrient regimes reflecting gradients typical of this region. Of these, seven showed P_i limitation, five revealed nitrogen (N) limitation, three exhibited light (L) limitation, and one bioassay had no growth. We found the phytoplankton community to shift from being predominately N limited in the early spring (March) to P limited in late spring and summer (May and July). Light limitation of phytoplankton production was recorded in several bioassays in July in water samples collected after peak annual flows from both the Mississippi and Atchafalaya Rivers. We also found that organic phosphorus, as glucose-6-phosphate, alleviated P limitation while phosphono-acetic acid had no effect. Whereas DIN/P_i and DIN/Si ratios in the initial water samples were good predictors of the outcome of phytoplankton production in response to inorganic nutrients, alkaline phosphatase activity was the best predictor when examining organic forms of phosphorus. We measured the rates of integrated primary production (0.33?C7.01 g C m^?2 d^?1), finding the highest rates within the Mississippi River delta and across Atchafalaya Bay at intermediate salinities. The lowest rates were measured along the outer shelf at the highest salinities and lowest nutrient concentrations (<0.1 ??M DIN and Pi). The results of this study indicate that P_i limitation of phytoplankton delays the assimilation of riverine DIN in the summer as the plume spreads across the shelf, pushing primary production over a larger region. Findings from water samples, taken adjacent the Atchafalaya River discharge, highlighted the importance of this riverine system to the overall production along the Louisiana coast.     

Nutrient enrichments and phytoplankton growth in the surface waters of the Louisiana Bight

Nitrate concentrations have increased twofold in the Mississippi River during the past three decades. The increased nitrogen loading to the Louisiana shelf has been postulated as a factor leading to eutrophication and the subsequent development of hypoxia west of the Mississippi River delta. While ratios of nitrogen:phosphorus and nitrogen:silica are relatively high in surface waters on the western Louisiana shelf, nitrogen has been posed as the ‘limiting’ nutrient in this region. Bioassays were performed with nutrient additions to surface waters collected from the Louisiana shelf to examine the potential for specific nutrient limitation. Experiments were conducted in March and September 1991, and May 1992. The growth responses of natural and cultured phytoplankton populations were determined by measuring the time course of in vivo and 3-(3,4 dichlorophenyl)-1, 1-dimethylurea (DCMU)-induced fluorescence, as well as initial and final chlorophylla concentrations. The results suggest that phosphate and silicate potentially limit phytoplankton growth during the winter-spring, particularly at low salinities. In late summer, in contrast, nitrogen limitation may be prominent at higher salinities.     

Phosphorus and nitrogen inputs to Florida Bay: The importance of the everglades watershed

A large environmental restoration project designed to improve the hydrological conditions of the Florida Everglades and increase freshwater flow to Florida Bay is underway. Here we explore how changing freshwater inflow to the southern Everglades is likely to change the input of nutrients to Florida Bay. We calculated annual inputs of water, total phosphorus (TP), total nitrogen (TN), and dissolved inorganic nitrogen (DIN) to Everglades National Park (ENP) since the early 1980s. We also examined changes in these nutrient concentrations along transects through the wetland to Florida Bay and the Gulf of Mexico. We found that the interannual variability of the water discharge into ENP greatly exceeded the interannual variability of flow-weighted mean nutrient concentrations in this water. Nutrient inputs to ENP were largely determined by discharge volume. These inputs were high in TN and low in TP; for two ENP watersheds TN averaged 1.5 mg l^?1 (0.11 mM) and 0.9 mg l^?1 (0.06 mM) and TP averaged 15 μg l^?1 (0.47 μM) and 9 μg l^?1 (0.28 μM). Both TP and DIN that flowed into ENP wetlands were rapidly removed from the water. Over a 3-km section of Taylor Slough, TP decreased from a flow-weighted mean of 11.6 μg l^?1 (0.37 μM) (0.20 μM) and DIN decreased from 240 μg l^?1 (17μM) to 36 μ l^?1 (2.6 μM). In contrast, TN, which was generally 95% organic N, changed little as it passed through the wetland. This resulted in molar TN:TP ratios exceeding 400 in the wetland. Decreases in TN concentrations only occurred in areas with relatively high P availability, such as the wetlands to the north of ENP and in the mangrove streams of western ENP. Increasing freshwater flow to Florida Bay in an effort to restore the Everglades and Florida Bay ecosystems is thus not likely to increase P inputs from the freshwater Everglades but is likely to increase TN inputs. Based on a nutrient budget of Florida Bay, both N and P inputs from the Gulf of Mexico greatly exceed inputs from the Everglades, as well as inputs from the atmosphere and the Florida Keys. We estimate that the freshwater Everglades contribute <3% of all P inputs and <12% of all N inputs to the bay. Evaluating the effect of ecosystem restoration efforts on Florida Bay requires greater understanding of the interactions of the bay with the Gulf of Mexico and adjacent mangrove ecosystems.     

Seasonal and spatial water quality changes in the outflow plume of the Atchafalaya River,Louisiana, USA

The objective of this study was to examine the interaction between the Atchafalaya River and the Atchafalaya Delta estuarine complex. Measurements of suspended sediments, inorganic nutrients (NO₃ ^?, NH₄ ⁺, PO₄ ^3?), chlorophylla (chla), and-salinity were taken monthly from December 1996 to January 1998. These data were compiled by season, and the Atchafalaya River plume data were also analyzed using the Generalized Additive Model technique. There were significant decreases in NO₃ ^? concentrations during summer, fall, and winter as river water passed through the estuary, that were attributable to chemical and biological processes rather than dilution with ambient water. In some regions there were higher chla concentrations during summer and fall compared to winter and spring, when river discharge and the introduction of inorganic nutrients were highest, suggesting biological processes were active during this study. The presence of NH₄ ⁺, as a percentage of available dissolved inorganic nitrogen, increased with distance from the Atchafalaya River, indicative of remineralization processes and NO₃ ^? reduction. Mean PO₄ ^3? concentrations were often higher in the estuarine regions compared to the Atchafalaya River. During summer total suspended solid (TSS) concentrations increased with distance from the river mouth, suggesting a turbidity maximum. Highest chla concentrations were found in the bayous and shallow water bodies of the Terrebonne marshes, as were the lowest TSS concentrations. The low chla concentrations found in other areas of this study, despite high inorganic nutrient concentrations, suggest light limitation as the major control of phytoplankton growth. Salinity reached near seawater concentrations at the outer edge of the Atchafalaya River plume, but much lower salinities (<10 psu) were observed at all other regions. The Atchafalaya Delta estuarine complex buffers the impact of the Atchafalaya River on the Louisiana coastal shelf zone, with a 41% of 47% decrease in Atchafalaya River NO₃ ^? concentrations before reaching Gulf waters.     

Mapping Phytoplankton Community Physiology on a River Impacted Continental Shelf: Testing a Multifaceted Approach

Seasonal phosphorus limitation occurs on the Louisiana continental shelf as a result of high nitrogen loads in the spring and early summer. Prior studies have assessed such nutrient limitation by laborious and time-consuming nutrient analyses, enzyme assays, and nutrient addition bioassays. We undertook surface (0.5–1 m) mapping of fast repetition rate fluorescence (FRRF) parameters to assess nutrient limitation in real time on the Louisiana continental shelf and Mississippi River plume from 29 June to 08 July, 2002 in an effort to further understand phytoplankton productivity in this region, as well as to better inform effective nutrient management strategies. Surface nutrient concentrations (NO₃⁻, NO₂⁻, NH₄⁺, PO₄³⁻), chlorophyll a biomass, alkaline phosphatase (AP) activity, and four FRRF parameters: the maximum quantum yield of photochemistry (F _v /F _m ), the functional absorption cross section for PSII, the time constant for Q _A reoxidation, and the connectivity factor, were measured during continuous underway mapping. Results from traditional methods to assess phytoplankton nutrient stress indicated widespread phosphorus limitation from the Mississippi River plume to the Atchafalaya River, manifested as high inorganic N/P ratios and elevated AP activities associated with phytoplankton biomass. The FRRF data indicated complex patterns of phytoplankton physiology that were likely driven by the rapidly changing conditions in local surface waters and heterogeneous phytoplankton community structure. Correlations of nutrient data and enzyme assays with FRRF parameters were significant but low, potentially due to differences in the manner and time scale with which nutrient limitation affects the different techniques used, indicating that further work is needed to interpret FRRF parameters in large, heterogeneous environments such as estuaries and continental shelves.     

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.     

Of manatees, mangroves, and the Mississippi River: Is there an estuarine signature for the Gulf of Mexico?

Important parameters of estuarine variability include morphology, flushing times, nutrient loading rates, and wetland: water ratios. This variability both reflects and disguises underlying relationships between the physics and biology of estuaries, which this comparative analysis seeks to reveal, using the Gulf of Mexico (GOM) estuaries as a starting point. A question used to focus this analysis is: are the GOM estuaries unique? The GOM receives the Mississippi River, a uniquely large, world-class river, which dominates the freshwater and nutrient inflows to the GOM continental shelf, whose margins include 35 major estuarine systems. These GOM estuaries have 28% and 41% of the U.S. estuarine wetlands and open water, respectively. Within the GOM, estuarine nitrogen, phosphorus, and suspended matter loading varies over 2 orders of magnitude. Anoxic estuarine events tend to occur in estuaries with relatively slow freshwater turnover and high nitrogen loading. Compared to estuaries from other regions in the U.S., the average GOM estuary is distinguished by shallower depths, faster freshwater flushing time, a higher wetland area:open water area ratio, greater fisheries yield per area wetland, lower tidal range, and higher sediment accumulation rates. The average GOM estuary often, but not always, has a flora and fauna not usually found in most other U.S. estuaries (e.g., manatees and mangroves). Coastal wetland loss in the GOM is extraordinarily high compared to other regions and is causally linked to cultural influences. Variations in nutrient loading and population density are very large among and within estuarine regions. This variation is large enough to demonstrate that there are insufficient systematic differences among these estuarine regions that precludes cross-system analyses. There are no abrupt discontinuities among regions in the fisheries yields per wetland area, tidal amplitude and vegetation range, salt marsh vertical accretion rates and organic accumulations, nitrogen retention, or wetland restoration rates. These results suggest that a comparative analysis emphasizing forcing functions, rather than geographic uniqueness, will lead to significant progress in understanding how all estuaries function, are perturbed, and even how they can be restored.     

Spatial and Temporal Patterns in <Emphasis Type="Italic">Thalassia testudinum</Emphasis> Leaf Tissue Nutrients at the Chandeleur Islands,Louisiana, USA

Seagrasses are submerged marine plants that are anchored to the substrate and are therefore limited to assimilating nutrients from the surrounding water column or sediment, or by translocating nutrients from adjacent shoots through the belowground rhizome. As a result, seagrasses have been used as reliable ecosystem indicators of surrounding nutrient conditions. The Chandeleur Islands are a chain of barrier islands in the northern Gulf of Mexico that support the only marine seagrass beds in Louisiana, USA, and are the sole location of the seagrass Thalassia testudinum across nearly 1000 km of the coastline from west Florida to central Texas. Over the past 150 years, the land area of the Chandeleur Islands has decreased by over half, resulting in a decline of seagrass cover. The goals of this study were to characterize the status of a climax seagrass species at the Chandeleur Islands, T. testudinum, in terms of leaf nutrient (nitrogen [N] and phosphorus [P]) changes over time, from 1998 to 2015, and to assess potential drivers of leaf nutrient content. Thalassia testudinum leaf nutrients displayed considerable interannual variability in N and P content and molar ratios, which broadly mimicked patterns in annual average dissolved nutrient concentrations in the lower Mississippi River. Hydrological modeling demonstrated the potential for multiple scenarios that would deliver Mississippi River water, and thus nutrients, to T. testudinum at the Chandeleur Islands. Although coastal eutrophication is generally accepted as the proximate cause for seagrass loss globally, there is little evidence that nutrient input from the Mississippi River has driven the dramatic declines observed in seagrasses at the Chandeleur Islands. Rather, seagrass cover along the Chandeleur Islands appears to be strongly influenced by island geomorphological processes. Although variable over time, the often elevated nutrient levels of the climax seagrass species, T. testudinum, which are potentially driven by river-derived nutrient inputs, raises an important consideration of the potential loss of the ecosystem functions and services associated with these declining seagrass meadows.     

Nutrient Limitation on Phytoplankton Growth in the Upper Barataria Basin,Louisiana: Microcosm Bioassays

The Davis Pond Diversion (DPD) was constructed to divert Mississippi River (MR) water into the Barataria Basin to reduce the salinity in support of wetland restoration on the Louisiana coast. To assess the phytoplankton nutrient limitation in adjacent water systems and potential impacts of DPD, 12 seasonal nutrient-phytoplankton bioassay experiments were conducted from October 2003 to July 2004 using the natural phytoplankton assemblages from freshwater and brackish-water lakes, Cataouatche and Salvador, LA (USA), which receive Mississippi River water from the DPD, and from a nearby freshwater lake, Lac des Allemands, that does not. Dissolved inorganic nitrogen (N), phosphorus (P), and silicate (Si) were added with different combinations at Redfield ratios in 10-l microcosms. Nitrogen was found to be the sole or primary limiting nutrient in all 12 experiments. N and P colimitations were found in seven of 12 experiments, but N was always the stronger limiting factor. P limitation was never observed to be the sole limiting nutrient. The results showed that a low concentration of P and a relatively high concentration of N do not necessarily indicate only P limitation in these lakes. Lake Cataouatche and Lake Salvador were dominated by centric diatoms, and Anabaena spp. were detected at high levels, particularly in summer. Lac des Allemands was generally dominated by N-fixing Anabaena spp. and other cyanobacteria, and their biomass responded significantly to N addition but not to P addition, indicating that nitrogen fixation in Lac des Allemands may be inhibited by other factors such as iron. Our bioassay results demonstrate that whether a water body is N- or P-limited is the consequence of the nutrient status and not the salinity regime. The results suggest that the addition of nutrient-rich waters via diversions of Mississippi River water into these lakes might increase the frequency of algal blooms, including noxious and toxic freshwater cyanobacteria.     

Inputs,transformations, and transport of nitrogen and phosphorus in Chesapeake Bay and selected tributaries

In this paper we assemble and analyze quantitative annual input-export budgets for total nitrogen (TN) and total phosphorus (TP) for Chesapeake Bay and three of its tributary estuaries (Potomac, Patuxent, and Choptank rivers). The budgets include estimates of TN and TP sources (point, diffuse, and atmospheric), internal losses (burial in sediments, fisheries yields, and denitrification), storages in the water column and sediments, internal cycling rates (zooplankton excretion and net sediment-water flux), and net downstream exchange. Annual terrestrial and atmospheric inputs (average of 1985 and 1986 data) of TN and TP ranged from 4.3 g TN m^?2 yr^?1 to 29.3 g TN m^?2 yr^?1 and 0.32 g TP m^?2 yr^?1 to 2.42 g TP m^?2 yr^?1, respectively. These rates of TN and TP input represent 6-fold to 8-fold and 13-fold to 24-fold increases in loads to these systems since the precolonial period. A recent 11-yr record for the Susquehanna River indicates that annual loads of TN and TP have varied by about 2-fold and 4-fold, respectively. TN inputs increased and TP inputs decreased during the 11-yr period. The relative importance of nutrient sources varied among these estuaries: point sources of nutrients delivered about half the annual TN and TP load to the Patuxent and nearly 60% of TP inputs to the Choptank; diffuse sources contributed 60–70% of the TN and TP inputs to the mainstream Chesapeake and Potomac River. The direct deposition of atmospheric wet-fall to the surface waters of these estuaries represented 12% or less of annual TN and TP loads except in the Choptank River (37% of TN and 20% of TP). We found direct, although damped, relationships between annual rates of nutrient input, water-column and sediment nutrient stocks, and nutrient losses via burial in sediments and denitrification. Our budgets indicate that the annual mass balance of TN and TP is maintained by a net landward exchange of TP and, with one exception (Choptank River), a net seaward transport of TN. The budgets for all systems revealed that inorganic nutrients entering these estuaries from terrestrial and atmospheric sources are rapidly converted to particulate and organic forms. Discrepancies between our budgets and others in the literature were resolved by the inclusion of sediments derived from shoreline erosion. The greatest potential for errors in our budgets can be attributed to the absence of or uncertainties in estimates of atmospheric dry-fall, contributions of nutrients via groundwater, and the sedimentation rates used to calculate nutrient burial rates.     

Seasonal Alternation Between Groundwater Discharge and Benthic Coupling as Nutrient Sources in a Shallow Coastal Lagoon

Submarine groundwater discharge (SGD) is now recognized as an important source of nutrients and freshwater to some coastal environments. We studied a shallow coastal lagoon (Little Lagoon, AL, USA) in the northern Gulf of Mexico that lacks riverine inputs but has been suspected to receive significant SGD. We observed persistent salinity gradients between the east and west ends of the lagoon and the pass connecting it to the Gulf of Mexico. Covariance between salinity in the lagoon and the groundwater tracer ²²²Rn indicated that SGD was responsible for the salinity gradients and is the primary source of freshwater to the lagoon. Cluster analysis of 246 biweekly samples based on temperature, salinity, and two proxies of SGD revealed two hydrographic regimes with different drivers for nutrient inputs. In samples characterized by high discharge and low temperatures (generally December–April), total nitrogen (TN) was negatively correlated with salinity, while total phosphorus (TP) was positively correlated with temperature. Total nitrogen in the groundwater was very high (0.36–4.80 mM) while total phosphorus was relatively low (0.3–2.3 μM), consistent with SGD as the source of TN during the high-discharge periods. In periods with low discharge and higher temperatures (approx. May–November), TN and TP had strong positive correlations with temperature and are inferred to originate from benthic efflux. Seasonal changes in nutrient stoichiometry in the lagoon water column also indicate an alternation between low TN/TP sediments and high TN/TP groundwater as the primary sources of nitrogen in this system.     

Comparison of nutrient and trace element distributions in the delta and shelf outflow regions of the Mississippi and Atchafalaya Rivers

Nutrient and trace element distributions were determined in the outflow region of the Mississippi and Atchafalaya rivers during high river discharge. This outflow region can be divided into two physiographic areas: the broad, shallow Louisiana Shelf off the Atchafalaya River and the narrow shelf off the Mississippi Delta. The physiographic differences between these two areas lead to observable differences in the chemical distributions. During high discharge conditions, nutrient depletion occurs at lower salinities on the Louisiana Shelf, relative to the delta outflow plume, and significant uptake of nickel and cadmium is apparent in these shelf waters, too. Important factors that appear to connect the physiographic to the chemical include the fate of the fluvial suspended load, rates of mixing, and the extent of productivity supported by recycled nutrients. The results suggest that the Mississippi-Atchafalaya outflow region may provide a natural laboratory for examining the possible effects of sea-level change on the biogeochemistry of estuarine and coastal environments.     

Differential salinity response between two Mississippi River subdeltas: Implications for changes in plant composition

Many studies have determined the physical dynamics of salt-water movement into riverine systems, but few studies have elucidated the role of periodic salinity incursions in affecting primary succession in active deltaic wetlands. The emergence of the neighboring Atchafalaya and Wax Lake Deltas, since a record Mississippi River flood in 1973, has created a unique area of land building and primary succession within the deteriorating wetland landscape of Louisiana. The vegetation in these deltas experiences the extremes of allogenic (riverine and tidal flooding, sedimentation, storm disturbance) and autogenic (herbivory, plant competition) forces. The rapid decline of the dominant vegetation,Sagittaria latifolia Willd. (Broadleaf Arrowhead), in the Atchafalaya Delta, and the continued dominance of this species in the Wax Lake Delta prompted us to investigate whether periodic salinity incursions were an additional stress on this species. The objectives of our study were to determine if salinity differences occurred between these deltas, describe the forces that controlled salinity incursions, and the level of salt exposure that adversely affected the growth ofS. latifolia. Continuous measurements of salinity and water level at six sites, revealed that salinity pulses (up to 7.0‰) were common in the Atchafalaya Delta; the Wax Lake Delta was not prone to elevated salinities. Salinity incursions in the Atchafalaya Delta were related to water level set-up, which was forced by strong (>10 m s⁻¹) easterly and southerly winds prior to cold front passages. The movement of distant tropical storms in the Gulf of Mexico produced salinity incursions of longer duration than cold fronts. In a greenhouse experiment, the growth ofS. latifolia was impaired by salinity concentrations of 6.0‰ within 13 d; flooding treatments (20 cm) compounded the loss of aboveground tissue by the end of the experiment. Results from the field and greenhouse studies, compared with historic evidence of elevated salinities in the Atchafalaya delta, indicate that periodic salinity incursions, combined with additional stresses such as flooding and vertebrate herbivory, influence the distribution or presence ofS. latifolia in the Atchafalaya Delta.     

Beneath the Salt Marsh Canopy: Loss of Soil Strength with Increasing Nutrient Loads

Although the broadly observed increase in nutrient loading rates to coastal waters in the last 100 years may increase aboveground biomass, it also tends to increase soil metabolism and lower root and rhizome biomass—responses that can compromise soil strength. Fourteen different multiyear field combinations of nutrient amendments to salt marshes were made to determine the relationship between soil strength and various nitrogen, phosphorus, and nitrogen+phosphorus loadings. There was a proportional decline in soil strength that reached 35% in the 60- to 100-cm soil layer at the highest loadings and did not level off. These loading rates are equivalent to those in the flow path of the Caernarvon river diversion, a major wetland restoration project near New Orleans; 12% of the wetlands in the flow path were converted to open water in 2005. The increased nutrient loading from the Mississippi River watershed this century has also driven the formation of the low oxygen zone (the “Dead Zone”) that forms off the Louisiana–Texas shelf each summer. These results suggest that improving water quality in the watershed will aid the restoration of both offshore waters and coastal wetland ecosystems.     

Characteristics of Danish estuaries

We review various aspects of the structure and functioning of Danish estuaries from data collected by the National Monitoring Program and from information in published sources. We present data on the physical, chemical, and biological characteristics of estuaries in Denmark, we evaluate the functioning of these systems as filters and transformers of nutrients and we evaluate the outlook for Danish estuaries in the future. Danish estuarine systems are for the most part shallow (<3 m deep), have short residence times, and tend to be heavily loaded with nutrients primarily from agricultural sources. Total average loads from land per unit watershed area are 112 kg P km^?2 yr^?1 and 2,400 kg N km^?2 yr^?1 during the period 1989–1995. The total phosphorus (TP) load in estuaries has been significantly reduced over the last decade, following implementation of the 1987 Action Plan for the Aquatic Environment (Vandmiljøplan in Danish) that prescribed that nitrogen loads to the total aquatic environment should be reduced by 50% and phosphorus loads by 80%. Reductions in the total nitrogen (TN) load have been more modest. Nutrient loading is one of the primary determinants of estuarine nutrient concentration with 70% of the annual variation in TN concentration and 55% of the annual variation in TP concentration explained by variation in the load. Many Danish estuaries have rich communities of macrophytes and benthic filter feeders, such asMytlis edulis andCiona intestinalis, that can control water column chlorophyll concentrations by their filter feeding activities. Many of the estuaries experience hypoxia and anoxia, especially during warm and calm summer months. Further reductions in nutrient loading are expected following implementation of the Action Plan for the Aquatic Environment II, with predicted improvements in oxygen concentrations and in the functioning of these shallow, dynamic estuarine systems.     

External nutrient sources,internal nutrient pools,and phytoplankton production in Chesapeake Bay

External nutrient loadings, internal nutrient pools, and phytoplankton production were examined for three major subsystems of the Chesapeake Bay Estuary—the upper Mainstem, the Patuxent Estuary, and the 01 Potomac Estuary—during 1985–1989. The atomic nitrogen to phosphorus ratios (TN:TP) of total loads to the 01 Mainstem, Patuxent, and the Potomac were 51, 29 and 35, respectively. Most of these loads entered at the head of the estuaries from riverine sources and major wastewater treatment plants. Approximately 7–16% for the nitrogen load entered the head of each estuary as particulate matter in contrast to 48–69% for phosphorus. This difference is hypothesized to favor a greater loss of phosphorus than nitrogen through sedimentation and burial. This process could be important in driving estuarine nitrogen to phosphorus ratios above those of inputs. Water column TN: TP ratios in the tidal fresh, oligohaline, and mesohaline salinity zones of each estuary ranged from 56 to 82 in the Mainstem, 27 to 48 in the Patuxent, and 72 to 126 in the Potomac. A major storm event in the Potomac watershed was shown to greatly increase the particulate fraction of nitrogen and phosphorus and lower the TN:TP in the river-borne loads. The load during the month that contained this storm (November 1985) accounted for 11% of the nitrogen and 31% of the phosphorus that was delivered to the estuary by the Potomac River during the entire 60-month period examined here. Within the Mainstem estuary, salinity dilution plots revealed strong net sources of ammonium and phosphate in the oligohaline to upper mesohaline region, indicating that these areas were sites of considerable internal recycling of nutrients to surface waters. The sedimentation of particulate nutrient loads in the upper reaches of the estuary is probably a major source of these recycled nutrients. A net sink of nitrate was indicated during summer. A combination of inputs and these internal recycling processes caused dissolved inorganic N to P ratios to approach 16:1 in the mesohaline zone of the Mainstem during late summer; this ratio was much higher at other times and in the lower salinity zones. Phytoplankton biomass in the mesohaline Mainstem reached a peak in spring and was relatively constant throughout the other seasons. Productivity was highest in spring and summer, accounting for approximately 33% and 44%, respectively, of the total annual productivity in this region. In the Patuxent and Potomac, the TN:TP ratios of external loads documented here are 2–4 times higher than those observed over the previous two decades. These changes are attributed to point-source phosphorus controls and the likelihood that nitrogen-rich nonpoint source inputs, including contributions from the atmosphere, have increased. These higher N:P ratios relative to Redfield proportions (16:1) now suggest a greater overall potential for phosphorus-limitation rather than nitrogen-limitation of phytoplankton in the areas studied.