Transport of groundwater-borne nutrients from watersheds and their effects on coastal waters

Anthropogenic activities on coastal watersheds increase nutrient concentrations of groundwater. As groundwater travels downslope it transports these nutrients toward the adjoining coastal water. The resulting nutrient loading rates can be significant because nutrient concentrations in coastal groundwaters may be several orders of magnitude greater than those of receiving coastal waters. Groundwater-borne nutrients are most subject to active biogeochemical transformations as they course through the upper 1 m or so of bottom sediments. There conditions favor anaerobic processes such as denitrification, as well as other mechanisms that either sequester or release nutrients. The relative importance of advective vs. regenerative pathways of nutrient supply may result in widely different rates of release of nutrients from sediments. The relative activity of denitrifiers also may alter the ratio of N to P released to overlying waters, and hence affect which nutrient limits growth of producers. The consequences of nutrient (particularly nitrate) loading include somewhat elevated nutrient concentrations in the watercolumn, increased growth of macroalgae and phytoplankton, reduction of seagrass beds, and reductions of the associated fauna. The decline in animals occurs because of habitat changes and because of the increased frequency of anoxic events prompted by the characteristically high respiration rates found in enriched waters.     

Eutrophication and macroalgal blooms in temperate and tropical coastal waters: nutrient enrichment experiments with Ulva spp.

Receiving coastal waters and estuaries are among the most nutrient‐enriched environments on earth, and one of the symptoms of the resulting eutrophication is the proliferation of opportunistic, fast‐growing marine seaweeds. Here, we used a widespread macroalga often involved in blooms, Ulva spp., to investigate how supply of nitrogen (N) and phosphorus (P), the two main potential growth‐limiting nutrients, influence macroalgal growth in temperate and tropical coastal waters ranging from low‐ to high‐nutrient supplies. We carried out N and P enrichment field experiments on Ulva spp. in seven coastal systems, with one of these systems represented by three different subestuaries, for a total of nine sites. We showed that rate of growth of Ulva spp. was directly correlated to annual dissolved inorganic nitrogen (DIN) concentrations, where growth increased with increasing DIN concentration. Internal N pools of macroalgal fronds were also linked to increased DIN supply, and algal growth rates were tightly coupled to these internal N pools. The increases in DIN appeared to be related to greater inputs of wastewater to these coastal waters as indicated by high δ¹⁵N signatures of the algae as DIN increased. N and P enrichment experiments showed that rate of macroalgal growth was controlled by supply of DIN where ambient DIN concentrations were low, and by P where DIN concentrations were higher, regardless of latitude or geographic setting. These results suggest that understanding the basis for macroalgal blooms, and management of these harmful phenomena, will require information as to nutrient sources, and actions to reduce supply of N and P in coastal waters concerned.     

Small Players, Large Role: Microbial Influence on Biogeochemical Processes in Pelagic Aquatic Ecosystems

Although prokaryotes are small in size, they are a significant biomass component in aquatic planktonic ecosystems and play a major role in biogeochemical processes. A review of the recent literature shows that the relative importance of prokaryotes to material and energy fluxes is maximized in low-productivity (oligotrophic) ecosystems and decreases in high-productivity (eutrophic) ecosystems. We conclude that competition with eukaryotic autotrophs for dissolved nutrients and competition with phagotrophic heterotrophs and physical processes (sinking, photooxidation) for organic carbon (C) play important roles in determining the relative abundance and impact of prokaryotes in aquatic systems. Oligotrophic systems have low nutrient concentrations, with high proportions of dissolved nutrients in organic form, which favors prokaryotic heterotrophs over phytoplankton. Furthermore, a high proportion of the available organic C is dissolved rather than particulate, which favors prokaryotic heterotrophs over phagotrophic heterotrophs. In eutrophic systems, increased relative concentrations and loading of inorganic nutrients and increased relative concentrations of particulate organic C select for phytoplankton and phagotrophic heterotrophs over prokaryotic heterotrophs. Increased particle sinking fluxes and/or decreased excretion of organic carbon (EOC) may also decrease the relative importance of prokaryotic heterotrophs in eutrophic systems. In oligotrophic systems, interactions between autotrophs and heterotrophs are tightly coupled because the dominant heterotrophs are similar in size and growth rates, as well as having similar nutrient composition to the dominant autotrophs, small phytoplankton. In eutrophic systems, increased productivity passes through zooplankton that are larger and have slower growth rates than the autotrophs, leading to a greater potential for decoupled auto- and heterotrophic production and increased export production. Received 18 July 2000; Accepted 13 September 2001.     

东太湖四角菱（Trapa quadrispinosa）生长特性

通过室外受控实验,研究了在不同沉积物类型及不同水体营养水平条件下东太湖四角菱的生长特性.结果表明:沉积物类型对菱的各项生长指标均有显著影响.在营养相对丰富的湖泥中,菱的相对生长率、同化根与吸收根的生物量以及株高增长率分别为相对贫瘠的岸泥的4.3、2.4、7.3、2.5倍,菱盘直径增长率在湖泥中为16.5%,在岸泥中则为负增长(-13.2%).水体营养盐浓度升高在一定程度上也促进了菱的生长,而且营养盐浓度(尤其是磷)升高对菱生长的促进作用在岸泥中要比在湖泥中显著得多.探讨了沉积物类型与水体营养水平对菱生长的作用机理,并认为水体富营养化的加剧及湖泊沼泽化所造成的氮、磷等营养物质在沉积物中的大量积累是东太湖菱种群迅速扩张的重要原因之一.     

沿岸海域富营养化与赤潮发生的关系

综述了赤潮的发生与沿岸海域富营养化的关系。近几十年来,人类活动使得天然水体的富营养化进程大大加速。营养负荷的增加与高生物量水华的增多相联系。控制营养输入后,浮游植物生物量或有害藻类水华事件也相应减少。营养的组成与浮游植物的种类组成及水华的形成有密切联系。有机营养对有害藻类水华的促进作用受到关注。营养输入时机影响浮游植物种间竞争的结果,因而对浮游植物的群落演替具有深远影响。由于浮游植物存在生理差异,因而对营养加富的反应因种而异。营养在调控某些有毒藻类的毒素产量方面也发挥着重要作用。此外,营养输入与藻类水华之间存在复杂的间接联系。当然,营养状况并非浮游植物群落演替的唯一决定因素。研究结果提示,控制营养输入、减缓水域富营养化是减少有害藻类水华发生的有效途径,而深入研究典型有害藻类的营养生理对策则为防治并最终消除有害藻类水华提供了理论基础。     

A feasibility analysis of discharge of non-contact,once-through industrial cooling water to forested wetlands for coastal restoration in Louisiana

Louisiana has had a high rate of coastal wetland loss due mainly to the isolation of the Mississippi River from the deltaic plain. We conducted a feasibility analysis of using once-through, non-contact industrial cooling water for restoring subsiding forested wetlands in coastal Louisiana. We considered the impacts of heated water and high nutrient and sediment concentrations. River diversions introduce sediments and nutrients to stimulate the productivity and accretion of coastal wetlands. Since increases in sediments and nutrients can cause water quality problems, we analyzed the assimilative capacity of the swamp. Based on a loading rates analysis, we estimated that the following nutrient reductions would occur: 75% for NO₃, 50% for TN, 60–75% for TP, and 100% for suspended sediments. Because of the concern of impacts from heated water, it is likely that the temperature of the cooling water will have to be decreased before discharge. Altering the duration and location of the discharge are ways to minimize the impact of temperature. We recommend that a pilot study be carried out to determine the effects of heated water on the functioning of the system, the retention of sediments and nutrients, and the impacts of different discharge scenarios.     

Impact of Submarine Groundwater Discharge on Marine Water Quality and Reef Biota of Maui

Generally unseen and infrequently measured, submarine groundwater discharge (SGD) can transport potentially large loads of nutrients and other land-based contaminants to coastal ecosystems. To examine this linkage we employed algal bioassays, benthic community analysis, and geochemical methods to examine water quality and community parameters of nearshore reefs adjacent to a variety of potential, land-based nutrient sources on Maui. Three common reef algae, Acanthophora spicifera, Hypnea musciformis, and Ulva spp. were collected and/or deployed at six locations with SGD. Algal tissue nitrogen (N) parameters (δ¹⁵N, N %, and C:N) were compared with nutrient and δ¹⁵N-nitrate values of coastal groundwater and nearshore surface water at all locations. Benthic community composition was estimated for ten 10-m transects per location. Reefs adjacent to sugarcane farms had the greatest abundance of macroalgae, low species diversity, and the highest concentrations of N in algal tissues, coastal groundwater, and marine surface waters compared to locations with low anthropogenic impact. Based on δ¹⁵N values of algal tissues, we estimate ca. 0.31 km² of Kahului Bay is impacted by effluent injected underground at the Kahului Wastewater Reclamation Facility (WRF); this region is barren of corals and almost entirely dominated by colonial zoanthids. Significant correlations among parameters of algal tissue N with adjacent surface and coastal groundwater N indicate that these bioassays provided a useful measure of nutrient source and loading. A conceptual model that uses Ulva spp. tissue δ¹⁵N and N % to identify potential N source(s) and relative N loading is proposed for Hawaiʻi. These results indicate that SGD can be a significant transport pathway for land-based nutrients with important biogeochemical and ecological implications in tropical, oceanic islands.     

Chesapeake Bay nutrient budgets — a reassessment

Recently published annual mass balances or budgets for nitrogen, phosphorus, and silicon in Chesapeake Bay have pictured the estuary as retaining a very large fraction, perhaps all, of the nutrients that enter from land drainage, the atmosphere, and anthropogenic discharges. However, these budgets have been based on estimates of the net exchanges of nutrients at the mouth of the bay or on the rates of accumulation of nutrients and sediments calculated from the distributions of various geochemical tracers in the sediments. While conceptually straightforward, the first approach is subject to large errors because it requires the determination of a small "signal" against a large background of tidal "noise". The second approach has led to overestimates of the nutrient trapping efficiency of the bay because tracer-derived sediment deposition rates have been multiplied by the surface area of the whole bay or various parts of the bay rather than by the smaller area of active sediment deposition. This approach is also incorrect because the average, long-term rates of sediment deposition measured by the geochemical tracers, including major floods, have been compared to shorter-term records of nutrient input.The more appropriate calculation of nutrient retention based on contemporaneous measurements of nutrient and sediment input and the chemical compositon of sediments accumulated in the estuary shows that Chesapeake Bay retains only some 3–6% of the nitrogen, 11–17% of the phosphorus and 33–83% of the silicon brought into its waters during a year in which no major flood occurred.This behavior suggests that current problems of estuarine eutrophication are more a consequence of present nutrient inputs than an inevitable or inescapable legacy of past enrichment. It also follows that the management or manipulation of nutrient loadings to esturies may lead to a more rapid response in environmental quality than previously predicted.     

NUTRIENTS AND THE PRODUCTIVITY OF ESTUARINE AND COASTAL MARINE ECOSYSTEMS

SUMMARY

Recent research on estuarine and coastal marine systems has revealed two particularly interesting things about nutrients and productivity. First is the observation that these areas are among the most intensively fertilized environments on earth. Second is the common finding that much of the characteristically high primary productivity of these shallow waters is supported by nutrients released or recycled by pelagic and benthic microheterotrophs. Since nutrient inputs to coastal areas have probably been increasing and are likely to continue to do so, it is particularly important to understand the relationship between nutrient loading and nutrient cycling and the extent to which their interactions may set the levels of primary and secondary production in coastal systems.

That some direct relationship exists between the input of nutrients and the productivity of higher trophic levels has been a principle of marine ecology since the turn of the century. It is surprisingly difficult, however, to find quantitative evidence showing that estuaries, lagoons, or other coastal waters respond to eutrophication by producing a larger biomass of animals. Part of this difficulty arises because the amount of nitrogen or phosphorus incorporated in animal tissue is a very small term in the total nutrient budget of an estuary, and the accuracy and precision of ecological field measurements may not be adequate to the task. In addition, the response of natural systems to nutrient enrichment is compounded by changes in climate, hydrography, harvesting effort and technology, and pollution.

Attempts to avoid some of these problems by carrying out controlled nutrient addition experiments in the field or with mesocosms have been much rarer in marine ecology than in limnology. The results that are available for such studies seem to suggest that there is a modest enhancement of primary production with nutrient addition, but that most of this extra organic matter is rapidly consumed, presumably by microheterotrophs. In other words, as nutrient inputs rise, so does the rate of nutrient recycling. Only a small fraction of the added nutrients appears as an increment in the production of higher trophic levels.     

Impact of nutrient loading on phytoplankton: a mesocosm experiment in the eutrophic Lake Taihu,China

Hydrobiologia - An increased nutrient loading drives eutrophication of lake ecosystems. Nutrient loading has two different origins: (1) internal loading due to nutrients release from sediments and...     

Effects of sunlight on the production of dissolved organic and inorganic nutrients from resuspended sediments

A series of laboratory-based and field experiments was conducted to address the effects of sunlight-exposed resuspended sediments on dissolved nutrient fluxes in two different water bodies. In suspensions of tidal creek sediments in 0.2 μm-filtered creek water, measurable increases in dissolved nutrients occurred after only 2 h of exposure to simulated sunlight. During a 6-h irradiation, nutrient release rates for total dissolved nitrogen (TDN) and phosphate were 2.2 ± 0.5 (standard error; S.E.) μmol g^?1 h^?1 and 0.09 ± 0.005 μmol g^?1 h^?1 (S.E.), compared to no significant changes in dark controls. The majority of nitrogen was released as dissolved organic nitrogen (87% on average) with lesser amounts of ammonium (13%). Continental shelf sediments resuspended in unfiltered seawater also released phosphate and TDN when exposed to sunlight, suggesting that this process can occur in a variety of marine and estuarine environments. The source material for inorganic nutrients appears to be associated with sediments rather than dissolved organic matter, as no significant changes in nutrient concentrations occurred in experiments with 0.2 μm-filtered creek water or seawater alone. Results suggest that photoproduction of dissolved nutrients from resuspended sediments could be an episodically significant and previously unrecognized source of dissolved organic and inorganic nutrients to coastal ecosystems. This process may be especially important for continental margins where episodic resuspension events occur, as well as in regions experiencing high riverine sediment fluxes resulting from erosion associated with deforestation and desertification.     

Using porewater profiles to assess nutrient availability in seagrass-vegetated carbonate sediments

We measured porewater profiles of inorganic (NH₄ ⁺, NO₃ ^–(+NO₂ ^–), PO₄ ^3– (hereafter referred to as DIP)) and organic (DON, DOP) nutrients in seagrass-vegetated sediments at two sites in a shallow bay in Bermuda within close proximity (200 m) but subject to different nutrient loading. At both sites, total dissolved and inorganic nutrient concentrations were usually 1–2 orders of magnitude higher in the sediments than in the water column, with the exception of NO₃ ^–. Organic N and P were significant components of the total dissolved nutrient pools both in the sediment porewater and in the overlying water column (up to 75% for DON and 40% for DOP), and may be important in meeting plant nutrient demands. We used two approaches to examine how well porewater nutrient concentrations reflected the relative availabilities of N and P for seagrasses: (1) a simple stoichiometric nutrient regeneration model based on the N:P ratio of decomposing organic matter and porewater NH₄ ⁺ concentrations to predict porewater DIP, and (2) fitting of the porewater profiles to estimate rates of net nutrient production (or consumption), which reflects the balance between nutrient sources and sinks in the rhizosphere. The stoichiometric model indicated that sediment porewaters were depleted in P relative to N in the low-nutrient outer bay site, and enriched in P relative to N in the higher-nutrient inner bay site. These results are consistent with the mechanism of carbonate sediments in oligotrophic tropical environments being a strong sink for dissolved inorganic P and our previous work suggesting that nutrient enrichment causes P to become disproportionately more available than N. Net nutrient production rates of porewater P at both sites and N at the inner bay site were low (typically < 2%) relative to the nutrient demands of the seagrasses. The implications of the profile interpretation are two-fold: (1) the low rates of net nutrient production indicate diffusive losses from the root zone were insignificant and that nutrient turnover rates were high, except in the P-limited outer bay where N accumulated in sediment porewaters; and (2) because standing stock nutrient concentrations often represent a small fraction of the total nutrients cycled in the sediments, they are in many cases a poor indicator of nutrient availability. Based on our estimates of losses from the root zone, decomposition, and plant uptake we have constructed a rough budget for the cycling of P and N at our two sites.     

Nitrogen cycling and anthropogenic impact in the tropical interamerican seas

We discuss the mechanisms leading to nutrient limitation in tropical marine systems, with particular emphasis on nitrogen cycling in Caribbean ecosystems. We then explore how accelerated nutrient cycling from human activities is affecting these systems.Both nitrogen and phosphorus exert substantial influence on biological productivity and structure of tropical marine ecosystems. Offshore planktonic communities are largely nitrogen limited while nearshore ecosystems are largely phosphorus limited. For phosphorus, the ability of sediment to adsorb and store phosphorus is probably greater for tropical carbonate sediments than for most nearshore sediments in temperate coastal systems. However, the ability of tropical carbonate sediments to take up phosphorus can become saturated as phosphorus loading from human sources increases. The nature of the sediment, the mixing rate between nutrient-laden runoff waters and nutrient-poor oceanic waters and the degree of interaction of these water masses with the sediment will probably control the dynamics of this transition.Nearshore tropical marine ecosystems function differently from their temperate counterparts where coupled nitrification/denitrification serves as an important mechanism for nitrogen depuration. In contrast, nearshore tropical ecosystems are more susceptible to nitrogen loading as depurative capacity of the microbial communities is limited by the fragility of the nitrification link. At the same time, accumulation of organic matter in nearshore carbonate sediments appears to impair their capacity for phosphorus immobilization. In the absence of depurative mechanisms for either phosphorus or nitrogen, limitation for both these nutrients is alleviated and continued nutrient loading fuels the proliferation of nuisance algae.     

Development and practical application of limiting values for the phosphate concentration in surface waters in the Netherlands

Summary To counteract eutrophication it is important to possess sufficient information (1) on the relationship between the content of nutrients and the algal biomass, and (2) on the cycling of nutrients in lakes.A comparative study of a large number of surface waters in the Netherlands has demonstrated that it is possible to derive a relationship between the nitrogen and phosphorus concentrations and the upper limit of the chlorophyll concentration, averaged over the summer season.For the authority in charge of water quality it is essential to know how far the phosphorus loading must be reduced to attain the desired phosphate concentration. The results of an extensive study of the nutrient budgets of Lake Wolderwijd-Nuldernauw over the period 1976–1979 demonstrate that in certain cases a relative high phosphate concentration can occur (approx. 0.30 mg/l) in spite of a low external phosphorus loading (approx. 0.7 g P/m² year). Such high concentrations can be explained by the continuous presence of a bloom of blue-green algae and the release of phosphate from the aquatic sediments during the summer. In such lakes it is necessary to take additional measures, such as flushing with water poor in algae and phosphates, or, where necessary, the removal of aquatic sediment rich in phosphate.     

Bacteria Contribute to Sediment Nutrient Release and Reflect Progressed Eutrophication-Driven Hypoxia in an Organic-Rich Continental Sea

In the sedimental organic matter of eutrophic continental seas, such as the largest dead zone in the world, the Baltic Sea, bacteria may directly participate in nutrient release by mineralizing organic matter or indirectly by altering the sediment’s ability to retain nutrients. Here, we present a case study of a hypoxic sea, which receives riverine nutrient loading and in which microbe-mediated vicious cycles of nutrients prevail. We showed that bacterial communities changed along the horizontal loading and vertical mineralization gradients in the Gulf of Finland of the Baltic Sea, using multivariate statistics of terminal restriction fragments and sediment chemical, spatial and other properties of the sampling sites. The change was mainly explained by concentrations of organic carbon, nitrogen and phosphorus, which showed strong positive correlation with Flavobacteria, Sphingobacteria, Alphaproteobacteria and Gammaproteobacteria. These bacteria predominated in the most organic-rich coastal surface sediments overlain by oxic bottom water, whereas sulphate-reducing bacteria, particularly the genus Desulfobacula, prevailed in the reduced organic-rich surface sediments in the open sea. They correlated positively with organic nitrogen and phosphorus, as well as manganese oxides. These relationships suggest that the bacterial groups participated in the aerobic and anaerobic degradation of organic matter and contributed to nutrient cycling. The high abundance of sulphate reducers in the surficial sediment layers reflects the persistence of eutrophication-induced hypoxia causing ecosystem-level changes in the Baltic Sea. The sulphate reducers began to decrease below depths of 20 cm, where members of the family Anaerolineaceae (phylum Chloroflexi) increased, possibly taking part in terminal mineralization processes. Our study provides valuable information on how organic loading affects sediment bacterial community compositions, which consequently may maintain active nutrient recycling. This information is needed to improve our understanding on nutrient cycling in shallow seas where the dead zones are continuously spreading worldwide.     

Sediment microbial enzyme activity as an indicator of nutrient limitation in Great Lakes coastal wetlands

1. We compared the extracellular enzyme activity (EEA) of sediment microbial assemblages with sediment and water chemistry, gradients in agricultural nutrient loading (derived from principal component analyses), atmospheric deposition and hydrological turnover time in coastal wetlands of the Laurentian Great Lakes. 2. There were distinct increases in nutrient concentrations in the water and in atmospheric N deposition along the gradient from Lake Superior to Lake Ontario, but few differences between lakes in sediment carbon (C), nitrogen (N) or phosphorus (P). Wetland water and sediment chemistry were correlated with the agricultural stress gradient, hydrological turnover time and atmospheric deposition. 3. The N : P ratio of wetland waters and sediments indicated that these coastal wetlands were N‐limited. Nutrient stoichiometry was correlated with the agricultural stress gradient, hydrological turnover time and atmospheric deposition. 4. Extracellular enzyme activity was correlated with wetland sediment and water chemistry and stoichiometry, atmospheric N deposition, the agricultural stress gradient and the hydrological turnover time. The ratios of glycosidases to peptidases and phosphatases yielded estimates of nutrient limitation that agreed with those based solely on nutrient chemistry. 5. This study, the first to link microbial enzyme activities to regional‐scale anthropogenic stressors, suggests that quantities and ratios of microbial enzymes are directly related to the concentrations and ratios of limiting nutrients, and may be sensitive indicators of nutrient dynamics in wetland ecosystems, but further work is needed to elucidate these relationships.     

The interactive effects of excess reactive nitrogen and climate change on aquatic ecosystems and water resources of the United States

Nearly all freshwaters and coastal zones of the US are degraded from inputs of excess reactive nitrogen (Nr), sources of which are runoff, atmospheric N deposition, and imported food and feed. Some major adverse effects include harmful algal blooms, hypoxia of fresh and coastal waters, ocean acidification, long-term harm to human health, and increased emissions of greenhouse gases. Nitrogen fluxes to coastal areas and emissions of nitrous oxide from waters have increased in response to N inputs. Denitrification and sedimentation of organic N to sediments are important processes that divert N from downstream transport. Aquatic ecosystems are particularly important denitrification hotspots. Carbon storage in sediments is enhanced by Nr, but whether carbon is permanently buried is unknown. The effect of climate change on N transport and processing in fresh and coastal waters will be felt most strongly through changes to the hydrologic cycle, whereas N loading is mostly climate-independent. Alterations in precipitation amount and dynamics will alter runoff, thereby influencing both rates of Nr inputs to aquatic ecosystems and groundwater and the water residence times that affect Nr removal within aquatic systems. Both infrastructure and climate change alter the landscape connectivity and hydrologic residence time that are essential to denitrification. While Nr inputs to and removal rates from aquatic systems are influenced by climate and management, reduction of N inputs from their source will be the most effective means to prevent or to minimize environmental and economic impacts of excess Nr to the nation’s water resources.     

Nutrient release and resuspension generated by ruffe (Gymnocephalus cernuus) and chironomids

1. Benthivorous fish may play an important role in internal nutrient loading. Ruffe are highly specialised, feeding exclusively on bottom animals; thus all nutrients released via their feeding are derived from the bottom and are new to the water column. The fish can also release nutrients from the sediment through resuspension while searching for food. 2. The aim of this study was to estimate experimentally in the laboratory the effect on water quality of resuspension and nutrient release by ruffe and bottom animals (chironomids). 3. Ruffe released nutrients during 8 h experiments as follows: total P 1.4, dissolved PO₄ 0.6, total N 24.0 and NH₄‐N 15.9 μg g^?1 WW h^?1. A decreasing trend in mass‐specific release was observed over time, probably because of starvation. The mass‐specific release of total N and NH₄‐N decreased as the mean weight of fish increased. The mean ratio of excreted N : P was 32. 4. In 26 h experiments with sediment and both ruffe and chironomids, ruffe increased nutrient concentrations and turbidity values significantly but chironomids had an effect only on turbidity. Neither ruffe nor chironomids affected the ratio of inorganic N : P concentrations. An interaction between ruffe and chironomids was found for turbidity. 5. According to these results, benthivorous fish may increase nutrient concentrations in the water column and need to be taken into account when estimating internal loading.     

Groundwater and pore water inputs to the coastal zone

Both terrestrial and marine forces drive underground fluid flows in the coastal zone. Hydraulic gradients on land result in groundwater seepage near shore and may contribute to flows further out on the shelf from confined aquifers. Marine processes such as tidal pumping and current-induced pressure gradients may induce interfacial fluid flow anywhere on the shelf where permeable sediments are present. The terrestrial and oceanic forces overlap spatially so measured fluid advection through coastal sediments may be a result of composite forcing. We thus define “submarine groundwater discharge” (SGD) as any and all flow of water on continental margins from the seabed to the coastal ocean, regardless of fluid composition or driving force. SGD is typically characterized by low specific flow rates that make detection and quantification difficult. However, because such flows occur over very large areas, the total flux is significant. Discharging fluids, whether derived from land or composed of re-circulated seawater, will react with sediment components. These reactions may increase substantially the concentrations of nutrients, carbon, and metals in the fluids. These fluids are thus a source of biogeochemically important constituents to the coastal ocean. Terrestrially-derived fluids represent a pathway for new material fluxes to the coastal zone. This may result in diffuse pollution in areas where contaminated groundwaters occur. This paper presents an historical context of SGD studies, defines the process in a form that is consistent with our current understanding of the driving forces as well as our assessment techniques, and reviews the estimated global fluxes and biogeochemical implications. We conclude that to fully characterize marine geochemical budgets, one must give due consideration to SGD. New methodologies, technologies, and modeling approaches are required to discriminate among the various forces that drive SGD and to evaluate these fluxes more precisely.     

GRACILARIA EDULIS (RHODOPHYTA) AS A BIOLOGICAL INDICATOR OF PULSED NUTRIENTS IN OLIGOTROPHIC WATERS

The response of the marine macroalga Gracilaria edulis (Gmelin) Silva to nutrient pulses of varying magnitude was investigated to test its applicability as a marine bioindicator at two oligotrophic locations. After exposure to nutrient pulses, algal amino acid, tissue nitrogen, and chlorophyll a content were assessed relative to algae incubated under control conditions (no nutrient enrichment). The smallest nutrient pulse involved a nutrient enrichment experiment conducted within a coral atoll, whereas two larger pulses resulted from sewage discharge to a tropical coastal bay. After exposure to the smallest nutrient pulse (10 × ambient), only changes in macroalgal amino acid concentration and composition were detected (mainly as increases in citrulline). At 100 × ambient concentrations, increases in tissue % nitrogen of the macroalgae were detected, in addition to responses in amino acids. Macroalgae exposed to the highest nutrient pulse (1000 × ambient) responded with increased chlorophyll a , tissue nitrogen, and amino acids within the three day incubation period. In contrast to these algal responses, analytical water sampling techniques failed to detect elevated nutrients when nutrient pulses were not occurring. The responses of this algal bioindicator to variable nutrient pulses may provide a useful tool for investigating the source and geographical extent of nutrients entering oligotrophic coastal waters.