National Science Education Standards

A primary focus of coastal science during the past 3 decades has been the question: How does anthropogenic nutrient enrichment cause change in the structure or function of nearshore coastal ecosystems? This theme of environmental science is recent, so our conceptual model of the coastal eutrophication problem continues to change rapidly In this review, I suggest that the early (Phase I) con- ceptual model was strongly influenced by limnologists, who began intense study of lake eutrophication by the 1960s The Phase I model emphasized changing nutrient input as a signal, and responses to that signal as increased phytoplankton biomass and primary production, decomposition of phytoplankton- derived organic matter, and enhanced depletion of oxygen from bottom waters Coastal research in recent decades has identified key differences in the responses of lakes and coastal-estuarine ecosystems to nutrient enrichment The contemporary (Phase II) conceptual model reflects those differences and includes explicit recognition of (1) system-specific attributes that act as a filter to modulate the responses to enrichment (leading to large differences among estuarine-coastal systems in their sensitivity to nu- trient enrichment); and (2) a complex suite of direct and indirect responses including linked changes in: water transparency, distribution of vascular plants and biomass of macroalgae, sediment biogeochem- istry and nutrient cycling, nutrient ratios and their regulation of phytoplankton community composition, frequency of toxic/harmful algal blooms, habitat quality for metazoans, reproduction/growth/survival of pelagic and benthic invertebrates, and subtle changes such as shifts in the seasonality of ecosystem functions Each aspect of the Phase II model is illustrated here with examples from coastal ecosystems around the world In the last section of this review I present one vision of the next (Phase III) stage in the evolution of our conceptual model, organized around 5 questions that will guide coastal science in the early 21st century: (1) How do system-specific attributes constrain or amplify the responses of coastal ecosystems to nutrient enrichment? (2) How does nutrient enrichment interact with other stressors (toxic contaminants, fishing harvest, aquaculture, nonindigenous species, habitat loss, climate change, hydro- logic manipulations) to change coastal ecosystems? (3) How are responses to multiple stressors linked? (4) How does human-induced change in the coastal zone impact the Earth system as habitat for humanity and other species? (5) How can a deeper scientific understanding of the coastal eutrophication problem be applied to develop tools for building strategies at ecosystem restoration or rehabilitation?

https://www.int-res.com/articles/meps/210/m210p223.pdf

Our evolving conceptual model of the coastal eutrophication problem

Denitrification is a critical process regulating the removal of bioavailable nitrogen (N) from natural and human-altered systems. While it has been extensively studied in terrestrial, freshwater, and marine systems, there has been limited communication among denitrification scientists working in these individual systems. Here, we compare rates of denitrification and controlling factors across a range of ecosystem types. We suggest that terrestrial, freshwater, and marine systems in which denitrification occurs can be organized along a continuum ranging from (1) those in which nitrification and denitrification are tightly coupled in space and time to (2) those in which nitrate production and denitrification are relatively decoupled. In aquatic ecosystems, N inputs influence denitrification rates whereas hydrology and geomorphology influence the proportion of N inputs that are denitrified. Relationships between denitrification and water residence time and N load are remarkably similar across lakes, river reaches, estuaries, and continental shelves. Spatially distributed global models of denitrification suggest that continental shelf sediments account for the largest portion (44%) of total global denitrification, followed by terrestrial soils (22%) and oceanic oxygen minimum zones (OMZs; 14%). Freshwater systems (groundwater, lakes, rivers) account for about 20% and estuaries 1% of total global denitrification. Denitrification of land-based N sources is distributed somewhat differently. Within watersheds, the amount of land-based N denitrified is generally highest in terrestrial soils, with progressively smaller amounts denitrified in groundwater, rivers, lakes and reservoirs, and estuaries. A number of regional exceptions to this general trend of decreasing denitrification in a downstream direction exist, including significant denitrification in continental shelves of N from terrestrial sources. Though terrestrial soils and groundwater are responsible for much denitrification at the watershed scale, per-area denitrification rates in soils and groundwater (kg Nkm � 2 � yr � 1 ) are, on average, approximately one-tenth the per-area rates of denitrification in lakes, rivers, estuaries, continental shelves, or OMZs. A number of potential approaches to increase denitrification on the landscape, and thus decrease N export to sensitive coastal systems exist. However, these have not generally been widely tested for their effectiveness at scales required to significantly reduce N export at the whole watershed scale.

/pdf/denitrification-across-landscapes-and-waterscapes-a-ei9e1947rb.pdf

Denitrification across landscapes and waterscapes: a synthesis.

The first special volume of Limnology and Oceanography, published in 1972, focused on whether phosphorus (P) or carbon (C) is the major agent causing eutrophication in aquatic ecosystems. Only slight mention was made that estuaries may behave differently from lakes and that nitrogen (N) may cause eutrophication in estuaries. In the following decade, an understanding of eutrophication in estuaries proceeded in relative isolation from the community of scientists studying lakes. National water quality policy in the United States was directed almost solely toward P control for both lakes and estuaries, and similarly, European nations tended to focus on P control in lakes. Although bioassay data indicated N control of eutrophication in estuaries as early as the 1970s, this body of knowledge was treated with skepticism by many freshwater scientists and water-quality managers, because bioassay data in lakes often did not properly indicate the importance of P relative to C in those ecosystems. Hence, the bioassay data in estuaries had little influence on water-quality management. Over the past two decades, a strong consensus has evolved among the scientific community that N is the primary cause of eutrophication in many coastal ecosystems. The development of this consensus was based in part on data from whole-ecosystem studies and on a growing body of evidence that presented convincing mechanistic reasons why the controls of eutrophication in lakes and coastal marine ecosystems may differ. Even though N is probably the major cause of eutrophication in most coastal systems in the temperate zone, optimal management of coastal eutrophication suggests controlling both N and P, in part because P can limit primary production in some systems. In addition, excess P in estuaries can interact with the availability of N and silica (Si) to adversely affect ecological structure. Reduction of P to upstream freshwater ecosystems can also benefit coastal marine ecosystems through mechanisms such as increased Si fluxes.

https://aslopubs.onlinelibrary.wiley.com/doi/pdf/10.4319/lo.2006.51.1_part_2.0364

Nitrogen as the limiting nutrient for eutrophication in coastal marine ecosystems: Evolving views over three decades

The second largest zone of coastal hypoxia (oxygen-depleted waters) in the world is found on the northern Gulf of Mexico continental shelf adjacent to the outflows of the Mississippi and Atchafalaya Rivers. The combination of high freshwater discharge, wind mixing, regional circulation, and summer warming controls the strength of stratification that goes through a well-defined seasonal cycle. The physical structure of the water column and high nutrient loads that enhance primary production lead to an annual formation of the hypoxic water mass that is dominant from spring through late summer. Paleoindicators in dated sediment cores indicate that hypoxic conditions likely began to appear around the turn of the last century and became more severe since the 1950s as the nitrate flux from the Mississippi River to the Gulf of Mexico tripled. Whereas increased nutrients enhance the production of some organisms, others are eliminated from water masses (they either emigrate from the area or die) where the...

Gulf of Mexico Hypoxia, A.K.A. “The Dead Zone”

Narragansett Bay is a relatively well-mixed, high salinity coastal embayment and estuary complex in southern New England (USA). Much of the shoreline is urban and the watershed is densely developed. We have combined our data on C, N, and P inputs to this system, on C, N, and P accumulation in the sediments, and on denitrification with extensive work by others to develop approximate annual mass balances for these elements. The results show that primary production within the bay is the major source of organic carbon (4 times greater than other sources), that land drainage and upstream sewage and fertilizer are the major sources of N, and that landward flowing bottom water from offshore may be a major source of dissolved inorganic phosphorus. Most of the nutrients entering the bay arrive in dissolved inorganic form, though DON is a significant component of the N carried by the rivers. About 40% of the DIN in the rivers is in the form of ammonia. Sedimentation rates are low in most of Narragansett Bay, and it appears that less than 20% of the total annual input of each of these elements is retained within the system. A very small amount of C, N, and P is removed in fisheries landings, denitrification in the sediments removes perhaps 10–25% of the N input, and most of the carbon fixed in the system is respired within it. Stoichiometric calculations suggest that some 10–20% of the organic matter formed in the bay is exported to offshore and that Narragansett Bay is an autotrophic system. Most of the N and P that enters the bay is, however, exported to offshore waters in dissolved inorganic form. This assessment of the overall biogeochemical behavior of C, N, and P in the bay is consistent with more rigorously constrained mass balances obtained using large living models or mesocosms of the bay at the Marine Ecosystem Research Laboratory (MERL).

An assessment of the annual mass balance of carbon, nitrogen, and phosphorus in Narragansett Bay

Nitrogen gas flux was measured from sediments taken from Narragansett Bay, Rhode Island, Boston Harbour, Massachusetts, and the Pawcatuck River Estuary, Rhode Island In addition to studies of field cores taken directly from these systems, intact sediments were taken from Narragansett Bay and maintained in control and nutrient enriched mesocosms Sediment denitrification was measured as a flux of N 2 gas from sediments in N 2 -free chambers The advantages of this technique are that it allows for the direct measurement of denitrification in undisturbed sediment cores under ambient conditions of dissolved nutrients, oxygen, and temperature The challenge of this technique has been to correctly distinguish between N 2 fluxes produced by denitrification activity and fluxes of N 2 caused by sediment porewater de-gassing In this study, anoxic 'control' cores were used to provide continuous checks on the magnitude of porewater de-gassing rates, and allowed measured rates of total N 2 flux to be corrected for this background flux The use of anoxic control cores allowed measurements to begin soon after core collection, without the need for long pre-incubations Observed rates of sediment denitrification ranged from 0 to 195 μmol N 2 m -2 h -1  Highest rates were found in the enriched mesocosms and at a sewage outfall site in Boston Harbour Denitrification rates increased exponentially with temperature and were enhanced by added nutrients Results from the anoxic control cores showed that even after 10 days, the background flux of N 2 from de-gassing porewaters (10-34 μmol N 2 m -2 h -1 ) remained a significant fraction of the overall N 2 flux Although rates of sediment denitrification were stimulated by added nutrients, the overall percentage loss of N from these systems through denitrification did not keep pace with N enrichment, generally accounting for less than 20% of N inputs Thus denitrification may not necessarily provide a significant pathway for alleviating eutrophication effects in coastal waters

The Effect of Temperature, Oxygen, Salinity, and Nutrient Enrichment on Estuarine Denitrification Rates Measured with a Modified Nitrogen Gas Flux Technique

The timing and magnitude of phytoplankton blooms have changed markedly in Narragansett Bay, RI (USA) over the last half century. The traditional winter–spring bloom has decreased or, in many years, disappeared. Relatively short, often intense, diatom blooms have become common in spring, summer, and fall replacing the summer flagellate blooms of the past. The annual and summer mean abundance (cell counts) and biomass (chl a ) of phytoplankton appear to have decreased based on almost 50 years of biweekly monitoring by others at a mid bay station. These changes have been related to warming of the water, especially during winter, and to increased cloudiness. A significant decline in the winter wind speed may also have played a role. The changes in the phenology of the phytoplankton and the oligotrophication of the bay appear to have decreased greatly the quantity and (perhaps) quality of the organic matter being deposited on the bottom of the bay. This decline has resulted in a very much reduced benthic metabolism as reflected in oxygen uptake, nutrient regeneration, and the magnitude and direction of the net flux of N 2 gas. Based on many decades of standard weekly trawls carried out by the Graduate School of Oceanography, the winter biomass of bottom feeding epibenthic animals has also declined sharply at the mid bay station. After decades of relatively constant anthropogenic nitrogen loading (and declining phosphorus loading), the fertilization of the bay will soon be reduced during May–October due to implementation of advanced wastewater treatment. This is intended to produce an oligotrophication of the urban Providence River estuary and the Upper Bay. The anticipated decline in the productivity of the upper bay region will probably decrease summer hypoxia in that area. However, it may have unanticipated consequences for secondary production in the mid and lower bay where climate-induced oligotrophication has already much weakened the historically strong benthic–pelagic coupling.

The impact of changing climate on phenology, productivity, and benthic–pelagic coupling in Narragansett Bay

Which nutrients limit primary production in coastal marine environments? Using large (13 000 1) mesocosms with sediments we explored nutrient limitation in Narragansett Bay, Rhode Island, USA, over a 9 wk period. Separate and combined additions of phosphorus and nitrogen were made to the enclosures. Phytoplankton biomass, daytime whole system oxygen production and nighttime whole system respiration showed an approximately 5-fold increase in nitrogen and nitrogen + phosphorus treatments, whereas phosphorus treatments had somewhat lower phytoplankton biomass and metabolism than controls. In these whole system experiments nitrogen was the nutrient most limiting to primary production.

/pdf/an-ecosystem-level-experiment-on-nutrient-limitation-in-oak0c3vk2s.pdf

An ecosystem level experiment on nutrient limitation in temperate coastal marine environments

In situ measurements of the exchange of ammonia, nitrate plus nitrite, phosphate, and dissolved organic phosphorus between sediments and the overlying water column were made in a shallow coastal lagoon on the ocean coast of Rhode Island, U.S.A. The release of ammonia from mud sediments in the dark (20–440 μmol per m2 per h) averaged ten times higher than from a sandy tidal flat (0–60 μmol per m2 per h), and while mud sediments also released nitrate and phosphate, sandy sediments took up these nutrients. Fluxes of nutrients from mud sediments, but not from sandy areas, markedly increased with temperature. Ammonia release rates for mud sediments in the light (0–350 μmol per m2 per h) were lower than those in the dark and it is estimated that some 25% of the ammonia released to the water column on an annual basis may be intercepted by the benthic microfloral community. Estimates of the annual net exchange of nutrients across the sediment-water interface, weighted by sediment type for the lagoon as a whole, showed a release of 450 mmol per m2 of ammonia, 5 mmol per m2 of phosphate, 5 mmol per m2 of dissolved organic phosphorus, and an uptake of 80 mmol per m2 of nitrate. Although rates of ammonia and nitrate exchange were comparable to those described for the deeper heterotrophic bottom communities of nearby Narragansett Bay, rates of benthic phosphate release were significantly lower. On an annual basis the Bay benthos released approximately 20 times more inorganic phosphate per unit area than did the lagoon benthos. As a result., the N/P ratio for the flux from the sediments was 74∶1 in the lagoon, compared with 16∶1 in “average” marine plankton and 8∶1 for the benthic flux from Narragansett Bay. The lack of remineralized phosphate in the lagoon, is reflected in water, column phosphate concentrations (always <1 μm) and water column N/P ratios (annual N/P=27) and suggests that the lagoon may show phosphate limitation rather than the nitrogen limitation commonly associated with marine systems.

Barbara L. Nowicki

Papers

An assessment of the annual mass balance of carbon, nitrogen, and phosphorus in Narragansett Bay

The Effect of Temperature, Oxygen, Salinity, and Nutrient Enrichment on Estuarine Denitrification Rates Measured with a Modified Nitrogen Gas Flux Technique

The impact of changing climate on phenology, productivity, and benthic–pelagic coupling in Narragansett Bay

An ecosystem level experiment on nutrient limitation in temperate coastal marine environments

Benthic nutrient remineralization in a coastal lagoon ecosystem