Response of water column microbial communities to sudden exposure to deltamethrin in aquatic mesocosms

Sudden exposure of an aquatic system to an insecticide can have significant effects on populations other than susceptible organisms. Although this is intuitively obvious, little is actually known about how such exposure might affect bacterial communities and their relative metabolic activity in ecosystems. Here, we assessed small sub-unit (ssu)-RNA levels in open and shaded 9 m(3) aquatic mesocosms (16 units - 2 x 2 factorial design in quadruplicate) to examine the effects of sudden addition of deltamethrin to the units. When deltamethrin was added, a cascade of bacterial then phytoplankton "blooms" occurred over time. The bacterial bloom, which most likely included organisms from the plastid/cyanobacterial phylogenetic guild, was almost immediate (within hours), whereas the phytoplankton (algal) bloom lagged by about 4 days. This sequential response can be explained by an apparent sudden release of nutrients consequent to arthropod death that triggered a series of responses in the microbial loop. Interestingly, bacterial blooms were noted in both open and shaded mesocosms, whereas the algal bloom was only seen in open units, suggesting that both deltamethrin addition (and presumptive nutrient release) and an adequate light supply was required for the phytoplankton response. Overall, this work shows that microbial activities as reflected by ssu-rRNA levels can respond dramatically via apparently indirect effects following insecticide application.     

Bottom-up excitable models of phytoplankton blooms

A simple nutrient-phytoplankton model is used to explore the dynamics of phytoplankton blooms. The model exhibits excitable behaviour in the sense that a large scale outbreak can only be triggered when a critical nutrient threshold is exceeded. The model takes into account several features often neglected but whose combined effect proves very important: (i) rapid nutrient recycling associated with the microbial loop and patch formation; (ii) self-shading; and (iii) a bottom-up approach, whereby nutrient levels are responsible for both the triggering and the demise of the bloom. Although the literature is replete with studies of ‘top-down’ models in which zooplankton grazing control the triggering and demise of the bloom, bottom-up models are nevertheless appropriate in many circumstances. We provide a full mathematical investigation of the effects of these three different features in an excitable system framework.     

160 Copepodology for the Phycologist with Apologies to G. E. Hutchenson

Heterocapsa triquetra is one of the most common bloom forming dinoflagellates found in estuaries and near shore regions around the world. In order to bloom, H. triquetra optimizes a suite of factors including low grazing pressure, increased nutrient inputs, alternative nutrient sources, and favorable salinity and hydrodynamic conditions, as well as the negative factors of temperature-limited growth, short day lengths, and periods of transient light limitation. The prevailing environmental conditions associated its wintertime blooms are largely the result of atmospheric forcing. Low-pressure systems moved through coastal area at frequent intervals and are accompanied by low air temperatures and rainfall. Runoff following the rainfall events supplies nutrients critical for bloom initiation and development. Heterocapsa triquetra blooms can reach chl a levels >100 mg L⁻¹ and cell densities between 1 to 6×106 L⁻¹. As the blooms develop, nutrient inputs from the river became insufficient to meet growth demand and H. triquetra feeds mixotrophically, reducing competition from co-occurring phytoplankton. Cloud cover associated with the low-pressure systems light limit H. triquetra growth as do low temperatures. More importantly though, low temperatures limit micro and macrozooplankton populations to such an extent that grazing losses are minimal.     

Comparing diatom and Alexandrium catenella/tamarense blooms in Thau lagoon: Importance of dissolved organic nitrogen in seasonally N-limited systems

Diatom blooms in Thau lagoon are always related to rain events leading to inputs of inorganic nutrients such as phosphate, ammonium and nitrate through the watershed with time lags of about 1 week. In contrast, blooms of Alexandrium catenella/tamarense can occur following periods of 3 weeks without precipitation and no significant input of conventional nutrients such as nitrate and phosphate. Field results also indicate a significant drop (from 22–25 to 15–16 μM over 3 days) in dissolved organic nitrogen (DON) at the bloom peak, as well as a significant inverse relationship between A. catenella/tamarense cell density and DON concentrations that is not apparent for diatom blooms. Such dinoflagellate blooms are also associated with elevated (6–9 μM) ammonium concentrations, a curious feature also observed by other investigators, possibly the results of ammonium excretion by this organism during urea or other organic nitrogen assimilation.The potential use of DON by this organism represents short cuts in the nitrogen cycle between plants and nutrients and requires a new model for phytoplankton growth that is different from the classical diatom bloom model. In contrast to such diatom blooms that are due to conventional (nitrate, phosphate) nutrient pulses, Alexandrium catenella/tamarense blooms on the monthly time scale are due to organic nutrient enrichment, a feature that allows net growth rates of about 1.3 d⁻¹, a value higher than that generally attributed to such organisms.     

Background nutrient concentration determines phytoplankton bloom response to marine heatwaves

Ocean temperature extreme events such as marine heatwaves are expected to intensify in coming decades due to anthropogenic global warming. Reported ecological and economic impacts of marine heatwaves include coral bleaching, local extinction of mangrove and kelp forests and elevated mortalities of invertebrates, fishes, seabirds and marine mammals. In contrast, little is known about the impacts of marine heatwaves on microbes that regulate biogeochemical processes in the ocean. Here we analyse the daily output of a near‐global ocean physical–biogeochemical model simulation to characterize the impacts of marine heatwaves on phytoplankton blooms in 23 tropical and temperate oceanographic regions from 1992 to 2014. The results reveal regionally coherent anomalies of shallower surface mixing layers and lower surface nitrate concentrations during marine heatwaves. These anomalies exert counteracting effects on phytoplankton growth through light and nutrient limitation. Consequently, the responses of phytoplankton blooms are mixed, but can be related to the background nutrient conditions of the study regions. The blooms are weaker during marine heatwaves in nutrient‐poor waters, whereas in nutrient‐rich waters, the heatwave blooms are stronger. The corresponding analyses of sea‐surface temperature, chlorophyll a and nitrate based on satellite observations and in situ climatology support this relationship between phytoplankton bloom anomalies and background nitrate concentration. Given that nutrient‐poor waters are projected to expand globally in the 21st century, this study suggests increased occurrence of weaker blooms during marine heatwaves in coming decades, with implications for higher trophic levels and biogeochemical cycling of key elements.     

Complexity in the eutrophication–harmful algal bloom relationship,with comment on the importance of grazing

This article seeks to guide the conceptual development and field application of the eutrophication–HAB hypothesis. After considering the evidence for this hypothesis, the importance of exogenous nutrients as a chemical habitat conditioner and the “family” of nutrient regulated effects that occur are discussed. The various definitions of eutrophication are applied; the conceptual ambiguity over how to perceive eutrophication, and the need to view eutrophication as a process and not as an ecological state are considered. The habitat irradiance-nutrient-flushing gradient regulates the bloom potential in response to exogenous nutrients. There is an apparent species-specific paradox within the eutrophication–HAB hypothesis related to the phycotoxin synthesis–nutrient relationship. Two nutrient–toxin relationships occur: toxin biosynthesis during nutrient sufficiency, and biosynthesis that requires a nutrient limitation. HAB events can be just as much nutrient depletion events as nutrient stimulated events, and whether the former develops depends upon the specific cellular toxicity–nutrient relationship of the bloom species. The importance of grazing in bloom regulation is highlighted. HABs and red tides generally should be viewed as blooms that are regulated by coupled nutrient-grazer processes – nutrient stimulation alone is inadequate, even when exogenous nutrients are not a factor. In assessing the eutrophication–HAB relationship, the collective grazing behavior of the micro-zooplankton, herbivorous copepods, filter feeding benthos, benthic larvae and, when present, omnivorous nekton must be considered. The importance of grazing in the bloom behavior of HAB species is illustrated using field and experimental data during a 5-month brown tide in Narragansett Bay. A cascade in grazing pressure regulated this bloom, with the initial collapse and then restoration of grazing pressure progressing through 7 stages of collective grazing pressure by micro-zooplankton, herbivorous copepods, benthic larvae, benthic filter feeders, and lytic virus infection.     

Top-down and bottom-up factors in tidepool communities

Recent studies suggest that nutrient variation influences rocky intertidal community structure, however empirical evidence is rare. In the Gulf of Maine, tidepools that occur on seagull feeding roosts are potentially subjected to regular nutrient loading from seagull guano. The results of a survey conducted on Swan's Island, ME show that roost tidepools have very low macroinvertebrate and macroalgal diversity as well as very high phytoplankton biomass compared to non-roost tidepools. An experiment presented here tested basic food chain hypotheses in tidepool communities. These basic food chain models predict that in a tidepool with one trophic level (phytoplankton only), phytoplankton biomass will increase when nutrients are enriched. In contrast, these models predict that in two trophic level tidepools (phytoplankton and mussels) herbivory will prevent an increase in phytoplankton biomass when nutrients are enriched. A short term 2×2 factorially designed field experiment was used to test this basic conceptual model using herbivory by mussels and enrichment with nitrogen as the main effects. The results of this investigation are consistent with the predictions of basic food chain models, and indicate that over the short time interval of a few days, herbivory by mussels is sufficient to maintain low phytoplankton levels following enrichment with nitrogen. Experimental enrichment with phosphorus in this study had no effect on phytoplankton biomass. The results of this study suggest that periodic pulses of nitrogen into tidepools will have little effect on phytoplankton biomass when mussels are present and that longer-term chronic nitrogen influxes may be driving the patterns of community structure in tidepools occurring on roosts.     

Ecology of phytoplankton communities dominated by Aureococcus anophagefferens: the role of viruses, nutrients, and microzooplankton grazing

We investigated the impact of viruses, nutrient loading, and microzooplankon grazing on phytoplankton communities in two New York estuaries that hosted blooms of the brown tide alga Aureococcus anophagefferens during 2000 and 2002. The absence of a bloom at one location during 2002 allowed for the fortuitous comparison of a bloom and non-bloom year at the same location as well as a comparison of two sites experiencing bloom and non-bloom conditions during the same year. During the study, blooms were found at locations with high levels of dissolved organic nitrogen and lower nitrate concentrations compared to a non-bloom location. Experimental additions of inorganic nitrogen and phosphorus yielded growth rates within the total phytoplankton community which significantly exceeded control treatments in 83% of experiments, while A. anophagefferens experienced significantly increased growth during only 20% of experimental inorganic nutrient additions. Consistent with prior research, these results suggest brown tides are not caused by eutrophication, but instead are more likely to occur when sources of labile DOM are readily available. Microzooplankton grazing rates on the total phytoplankton community during a bloom were lower than grazing rates at a non-bloom site, and grazing rates on A. anophagefferens were lower than grazing rates on the total community on some dates, suggesting that reduced grazing mortality may also promote brown tides. Mean densities of viruses during blooms (3 × 10⁸ ml⁻¹) were elevated compared to most estuarine environments and were twice the levels found at a non-bloom site. Experimental enrichment of the natural viral densities yielded a significant increase in A. anophagefferens growth rates relative to control treatments when background levels of viruses were low (<1.7 × 10⁸ ml⁻¹), suggesting that viruses may promote bloom occurrence by regenerating DOM or altering the composition of microbial communities.     

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。     

Student's tutorial on bloom hypotheses in the context of phytoplankton annual cycles

Phytoplankton blooms are elements in repeating annual cycles of phytoplankton biomass and they have significant ecological and biogeochemical consequences. Temporal changes in phytoplankton biomass are governed by complex predator–prey interactions and physically driven variations in upper water column growth conditions (light, nutrient, and temperature). Understanding these dependencies is fundamental to assess future change in bloom frequency, duration, and magnitude and thus represents a quintessential challenge in global change biology. A variety of contrasting hypotheses have emerged in the literature to explain phytoplankton blooms, but over time the basic tenets of these hypotheses have become unclear. Here, we provide a “tutorial” on the development of these concepts and the fundamental elements distinguishing each hypothesis. The intent of this tutorial is to provide a useful background and set of tools for reading the bloom literature and to give some suggestions for future studies. Our tutorial is written for “students” at all stages of their career. We hope it is equally useful and interesting to those with only a cursory interest in blooms as those deeply immersed in the challenge of understanding the temporal dynamics of phytoplankton biomass and predicting its future change.     

Rising CO2 Levels Will Intensify Phytoplankton Blooms in Eutrophic and Hypertrophic Lakes

Harmful algal blooms threaten the water quality of many eutrophic and hypertrophic lakes and cause severe ecological and economic damage worldwide. Dense blooms often deplete the dissolved CO₂ concentration and raise pH. Yet, quantitative prediction of the feedbacks between phytoplankton growth, CO₂ drawdown and the inorganic carbon chemistry of aquatic ecosystems has received surprisingly little attention. Here, we develop a mathematical model to predict dynamic changes in dissolved inorganic carbon (DIC), pH and alkalinity during phytoplankton bloom development. We tested the model in chemostat experiments with the freshwater cyanobacterium Microcystis aeruginosa at different CO₂ levels. The experiments showed that dense blooms sequestered large amounts of atmospheric CO₂, not only by their own biomass production but also by inducing a high pH and alkalinity that enhanced the capacity for DIC storage in the system. We used the model to explore how phytoplankton blooms of eutrophic waters will respond to rising CO₂ levels. The model predicts that (1) dense phytoplankton blooms in low- and moderately alkaline waters can deplete the dissolved CO₂ concentration to limiting levels and raise the pH over a relatively wide range of atmospheric CO₂ conditions, (2) rising atmospheric CO₂ levels will enhance phytoplankton blooms in low- and moderately alkaline waters with high nutrient loads, and (3) above some threshold, rising atmospheric CO₂ will alleviate phytoplankton blooms from carbon limitation, resulting in less intense CO₂ depletion and a lesser increase in pH. Sensitivity analysis indicated that the model predictions were qualitatively robust. Quantitatively, the predictions were sensitive to variation in lake depth, DIC input and CO₂ gas transfer across the air-water interface, but relatively robust to variation in the carbon uptake mechanisms of phytoplankton. In total, these findings warn that rising CO₂ levels may result in a marked intensification of phytoplankton blooms in eutrophic and hypertrophic waters.     

Role of toxin and nutrient for the occurrence and termination of plankton bloom--results drawn from field observations and a mathematical model

The termination of harmful algal blooms (HABs) and coexistence of phytoplankton-zooplankton populations are of great importance to human health, ecosystem, environment, tourism and fisheries. In this paper we propose a three-component model consisting of dissolved limiting nutrients (N) supplied at constant rate and partially recycled after the death of plankton by bacterial decomposition, phytoplankton (P) and zooplankton (Z), where the growth of zooplankton species reduce due to toxic chemicals released by phytoplankton species. Our analysis leads to different thresholds which are expressible in terms of model parameters and determine the existence and stability of various states of the system. We observe that phytoplankton-zooplankton persist if the maximal zooplankton ingestion rate exceeds a lower threshold value. It is shown that the coexistence equilibrium loses its stability when the dilution rate of the nutrient concentration passes through a critical value and Hopf bifurcation occurs that induces oscillations of the population. Our results indicate that the occurrence of bloom increases when the nutrient concentration is very high, and in that case toxin produced by the phytoplankton plays a very crucial role towards the termination of the planktonic bloom.     

Long-term nutrient trends and harmful cyanobacterial bloom potential in hypertrophic Lake Taihu,China

Rapid economic development in China’s Lake Taihu basin during the past four decades has accelerated nitrogen (N) and phosphorus (P) loadings to the lake. This has caused a shift from mesotrophic to hypertrophic conditions, symptomized by harmful cyanobacterial blooms (CyanoHABs). The relationships between phytoplankton biomass as chlorophyll a (Chla) and nutrients as total nitrogen (TN) and total phosphorus (TP) were analyzed using historical data from 1992 to 2012 to link the response of CyanoHAB potential to long-term nutrient changes. Over the twenty year study period, annual mean Chla showed significantly positive correlations with both annual mean TN and TP (P < 0.001), reflecting a strong phytoplankton biomass response to changes in nutrient inputs to the lake. However, phytoplankton biomass responded slowly to annual changes in TN after 2002. There was not a well-defined or significant relationship between spring TN and summertime Chla. The loss of a significant fraction of spring N loading due to denitrification likely weakened this relationship. Bioavailability of both N and P during the summer plays a key role in sustaining cyanobacterial blooms. The frequency of occurrence of bloom level Chla (>20 μg L^?1) was compared to TN and TP to determine nutrient-bloom thresholds. A decline in bloom risk is expected if TN remains below 1.0 mg L^?1 and TP below 0.08 mg L^?1.     

Application of an Algal Bloom Model (BLOOM II) to combat eutrophication

Summary The objective of the Algal Bloom Model (BLOOM II) is to predict the highest steady state value of the total biomass of all phytoplankton species in specified circumstances. Its calculations are usually on a weekly basis. Linear programming is being used to calculate the bloom, that may be constrained by the amounts of three nutrients (nitrogen, phosphorus and silicon) and light.Given a set of environmental conditions, the model will choose the optimal species composition among all species in the model (presently more than 10).The model has been calibrated for 3 model reservoirs and has been applied to about 15 different lakes in the Netherlands with highly different nutrient concentrations, background extinctions and mixing depths (from 1.2 to 15.0 m). Calculated biomass maxima, converted to microgram chlorophyll per litre vary from 50 to 600 and are remarkably close to the observations. Generally the model indicates the limiting factor(s) correctly; often also the species composition is correct.BLOOM II has been used to simulate the effects of changes in nutrient concentrations, mixing depths and flushing rates, among others, on the size, composition and time of the phytoplankton blooms, but so far little data are available to validate these predictions.     

东海原甲藻与中肋骨条藻的种间竞争数值模拟

研究探讨了两个零维箱式模型在东海典型赤潮藻东海原甲藻和中肋骨条藻竞争与演替研究中的应用。模型在采用不同接种密度下的单种培养实验数据进行参数校正后,被用来模拟不同N/P条件下单种培养实验以及两藻种共培养竞争实验,并以实验数据对其结果进行了验证。模拟结果表明,在单种培养条件下,模型能够较好地重现两种藻在不同N/P环境中的生长及对营养盐的利用;共培养实验的模拟结果显示,在所有初始细胞密度比例条件下,中肋骨条藻的最终密度均会超过东海原甲藻,且PO4的消耗主要源于中肋骨条藻的利用,与实验结果一致,表明模型能够很好地体现两种藻的竞争结果及对营养盐的竞争关系;由于模型不足以模拟除营养盐竞争以外的藻间相互作用,模拟结果未体现东海原甲藻细胞数迅速衰减这一现象,有待进一步研究。     

Dynamics of the ‘echo’ effect in a phytoplankton system with nitrogen fixation

Consideration of nitrogen fixation adds a positive nonlinear feedback to plankton ecosystem models. We investigate the consequences of this feedback for secondary phytoplankton blooms and the response of phytoplankton dynamics to physical forcing. The dynamics of phytoplankton, Trichodesmium (the nitrogen fixer), and nutrients is modeled with a system of three differential equations. The model includes two types of nonlinear interactions: the competition of phytoplankton and Trichodesmium for light, and the positive feedback resulting from Trichodesmium recycling. A typical simulation of the model in time, with forcing by a varying mixed-layer depth, reveals a clear successional sequence including a secondary or ‘echo’ bloom of the phytoplankton. We explain this sequence of events through the stability analysis of three different steady states of the model. Our analysis shows the existence of a critical biological parameter, the ratio of normalized growth rates, determining the occurrence of ‘echo’ blooms and the specific sequence of events following a physical perturbation. The interplay of positive and negative feedbacks appears essential to the timing and the type of events following such a perturbation.     

Phytoplankton growth rates in the freshwater tidal reaches of the Schelde estuary (Belgium) estimated using a simple light-limited primary production model

During the course of 1996, phytoplankton was monitored in the turbid, freshwater tidal reaches of the Schelde estuary. Using a simple light-limited primary production model, phytoplankton growth rates were estimated to evaluate whether phytoplankton could attain net positive growth rates and whether growth rates were high enough for a bloom to develop. Two phytoplankton blooms were observed in the freshwater tidal reaches. The first bloom occurred in March and was mainly situated in the most upstream reaches of the freshwater tidal zone, suggesting that it was imported from the tributary river Schelde. The second bloom occurred in July and August. This summer bloom was situated more downstream in the freshwater tidal reaches and appeared to have developed within the estuary. A comparison between phytoplankton growth rates estimated using a simple primary production model and flushing rate of the water indicated that no net increase in phytoplankton biomass was possible in March while phytoplankton could theoretically increase its biomass by 20% per day during summer. Chlorophyllaconcentrations at all times decreased strongly at salinities between 5–10 psu. This decline was ascribed to a combination of salinity stress and light limitation. Phytoplankton biomass and estimated annual net production were much higher in the freshwater tidal zone compared to the brackish reaches of the estuary (salinity > 10 psu) despite mixing depth to euphotic depth ratios being similar. Possible reasons for this high production include high nutrient concentrations, low zooplankton grazing pressure and import of phytoplankton blooms from the tributary rivers.     

How does eutrophication affect the role of grazers in harmful algal bloom dynamics?

Population dynamics of harmful algal bloom species are regulated both from the “bottom-up” by factors that affect their growth rate and from the “top-down” by factors that affect their loss rates. While it might seem apparent that eutrophication would have the greatest impact on factors affecting growth rates of phytoplankton (nutrient supply, light availability) the roles of top-down controls, including grazers and pathogens, cannot be ignored in studies of harmful bloom dynamics. Lags between the growth of phytoplankton and zooplankton populations, or disruption of zooplankton populations by adverse environmental conditions may be important factors in the initiation of plankton blooms under eutrophic conditions. Grazers that avoid feeding on harmful species and actively graze on competing species may also play important roles in bloom initiation. Grazers that are not affected by phytoplankton toxins and have growth rates comparable to phytoplankton (e.g. protozoan grazers) may have the potential to control the initiation of blooms. If the inhibition of grazers varies with cell density for blooms of toxic phytoplankton, eutrophication may increase the chances of blooms reaching threshold densities for grazer inhibition. In addition, secondary effects of eutrophication, including hypoxia and change in pH may adversely affect grazer populations, and further release HAB species from top-down control. The Texas brown tide (Aureoumbra lagunensis) blooms provide evidence for the role of grazer disruption in bloom initiation and the importance of high densities of brown tide cells in continued suppression of grazers.     

Positive feedbacks between bottom-up and top-down controls promote the formation and toxicity of ecosystem disruptive algal blooms: A modeling study

Harmful algal blooms that disrupt and degrade ecosystems (ecosystem disruptive algal blooms, EDABs) are occurring with greater frequency and severity with eutrophication and other adverse anthropogenic alterations of coastal systems. EDAB events have been hypothesized to be caused by positive feedback interactions involving differential growth of competing algal species, low grazing mortality rates on EDAB species, and resulting decreases in nutrient inputs from grazer-mediated nutrient cycling as the EDAB event progresses. Here we develop a stoichiometric nutrient–phytoplankton–zooplankton (NPZ) model to test a conceptual positive feedback mechanism linked to increased cell toxicity and resultant decreases in grazing mortality rates in EDAB species under nutrient limitation of growth rate. As our model EDAB alga, we chose the slow-growing, toxic dinoflagellate Karenia brevis, whose toxin levels have been shown to increase with nutrient (nitrogen) limitation of specific growth rate. This species was competed with two high-nutrient adapted, faster-growing diatoms (Thalassiosira pseudonana and Thalassiosira weissflogii) using recently published data for relationships among nutrient (ammonium) concentration, carbon normalized ammonium uptake rates, cellular nitrogen:carbon (N:C) ratios, and specific growth rate. The model results support the proposed positive feedback mechanism for EDAB formation and toxicity. In all cases the toxic bloom was preceded by one or more pre-blooms of fast-growing diatoms, which drew dissolved nutrients to low growth rate-limiting levels, and stimulated the population growth of zooplankton grazers. Low specific grazing rates on the toxic, nutrient-limited EDAB species then promoted the population growth of this species, which further decreased grazing rates, grazing-linked nutrient recycling, nutrient concentrations, and algal specific growth rates. The nutrient limitation of growth rate further increased toxin concentrations in the EDAB algae, which further decreased grazing-linked nutrient recycling rates and nutrient concentrations, and caused an even greater nutrient limitation of growth rate and even higher toxin levels in the EDAB algae. This chain of interactions represented a positive feedback that resulted in the formation of a high-biomass toxic bloom, with low, nutrient-limited specific growth rates and associated high cellular C:N and toxin:C ratios. Together the elevated C:N and toxin:C ratios in the EDAB algae resulted in very high bloom toxicity. The positive feedbacks and resulting bloom formation and toxicity were increased by long water residence times, which increased the relative importance of grazing-linked nutrient recycling to the overall supply of limiting nutrient (N).     

Nutrient limitation and algal blooms in urbanizing tidal creeks

Tidal creeks are commonly found in low energy systems on the East and Gulf Coasts of the United States, and are often subject to intense watershed human development. Many of these creeks are receiving urban and suburban runoff containing nutrients, among other pollutants. During the period 1993-2001, we studied three tidal creeks located in southeastern North Carolina, a rapidly urbanizing area. All three creeks received anthropogenic nutrient loading. Oligohaline to mesohaline stations in upper tidal creek regions had much higher nutrient (especially nitrate-N) concentrations than lower creek areas, and hosted spring and summer phytoplankton blooms that at times exceeded 200 μg chlorophyll a l⁻¹. Phytoplankton biomass during winter was low at all stations in all three creeks. Spring and summer nutrient addition bioassay experiments were conducted to characterize the nutrients limiting phytoplankton growth. Water from high salinity stations in all three creeks always showed significant positive responses to nitrate-N inputs, even at concentrations as low as 50 μg N l⁻¹. Low salinity stations in upper creek areas often showed significant responses to nitrate-N inputs, but on occasion showed sensitivity to phosphorus inputs as well, indicating the influence of anthropogenic nitrate loading. During several experiments, one of the upper stations showed no positive response to nutrient inputs, indicating that these stretches were nutrient replete, and further phytoplankton growth appeared to be light-limited either by phytoplankton self-shading or turbidity. Water from upper creek areas yielded much higher chlorophyll a concentrations in bioassay experiments than did lower creek water. In general, these urbanizing tidal creeks were shown to be very sensitive to nitrogen loading, and provide a physical environment conducive to phytoplankton bloom formation in nutrient-enriched areas. Tidal creeks are important ecological resources in that they are considered to be nursery areas for many species of fish and shellfish. To protect the ecological function of these small, but very abundant estuarine systems, management efforts should recognize their susceptibility to algal blooms and focus on control of nonpoint source nutrient inputs, especially nitrogen.