THE EFFECT OF NITROGEN SOURCE ON THE GROWTH AND TOXICITY OF THREE POTENTIALLY HARMFUL DINOFLAGELLATES

Increases in population and agriculture in coastal areas can result in increased nutrient inputs and alterations in the ratios of organic to inorganic nutrients in coastal waters. Such changes in coastal nutrient regimes can affect phytoplankton community structure by creating conditions favorable for growth and dominance of algae that were not dominant before. The effect that changes in ratios and concentrations of nutrients have on toxicity of harmful algal species is not well known. There seems to be a relationship; however, between nutrient stress and toxin production among harmful phytoplankton producing low‐N toxins, e.g. Diarrhetic Shellfish Poisoning (DSP) toxins. Even less is known about the relationship between organic nutrient uptake and toxin production. Benthic species and species in coastal areas are probably exposed to greater fluxes of dissolved organic nitrogen (DON). In this study, benthic and planktonic species of Prorocentrum were grown on L1 media with the sole N‐source varying among treatments as nitrate, ammonium, urea, L‐glutamic acid, and high molecular weight natural DON. An ELISA specific to the DSP toxins, okadaic acid and 35‐methylokadaic acid, was used to determine toxin production by each species when grown on the different N sources. Preliminary results indicate that some organic forms of N support growth as well as inorganic forms for Prorocentrum minimum, P. mexicanum, and P. hoffmannianum.     

<Emphasis Type="Italic">Prorocentrum rivalis</Emphasis> sp. nov. (Dinophyceae) and its phylogenetic affinities inferred from analysis of a mixed morphological and LSU rRNA data set

A new freshwater epiphytic Prorocentrum species, Prorocentrum rivalis, from the temperate region of the Haute-Vienne, France, is described. This species is the third freshwater species identified among approximately 60 marine Prorocentrum species. This new species is described using scanning electron microscope and phylogenetic analyses by a polyphasic approach (LSU rRNA sequences combined with 9 morphological characters). The phylogenetic analysis attests that P. rivalis is close to other planktonic freshwater species and the freshwater Prorocentrum clade is evolutionarily derived from an epiphytic freshwater prorocentroid ancestor. The unique marine species in the freshwater clade results from an ecophysiological reversion. P. rivalis differs from other epiphytic taxa by its rarity, its temperate distribution and its ecophysiological needs. The phylogeny confirms also that all planktonic Prorocentrum species are evolutionarily derived from epiphytic/benthic ancestors.     

Biological interactions in enclosed plankton communities including Alexandrium catenella and copepods: Role of phosphorus

A microcosm approach was used to test whether: a) growth under unbalanced nutrient conditions (varying N:P ratios) affected the susceptibility of a phytoplankton community including the dinoflagellate Alexandrium catenella (a paralytic shellfish toxin producer) to mesozooplankton grazing, and b) the potential effects of unbalanced nutrient conditions were mediated by changes in toxicity of A. catenella or by other mechanisms. The experimental setup consisted of fifteen 30 l microcosms, filled with water from the Barcelona Harbour and subjected to treatments combining nutrient inputs at three different N:P ratios (Redfield N:P ratio or nutrient-balanced, high N:P and low N:P), addition or omission of A. catenella (an estimated initial concentration of 38 A. catenella cells ml^− 1, a value typical for blooms in harbours of the Catalan coast), and selective addition of a cultured population of Acartia grani. P sufficiency had a strong positive effect on the growth of A. grani, both with or without A. catenella addition, presumably due to enhanced food quality of the prey community. The presence of this copepod resulted in lower concentrations of ciliates, A. catenella, and other dinoflagellates, suggesting active grazing by the copepods. No noxious effects of A. catenella on the copepods were detected at the relatively low cell concentrations of that dinoflagellate used in the experiment.     

Effects of nutrient-limiting supply ratios on toxin content of Karenia brevis grown in continuous culture

Toxins produced as secondary metabolites can play important roles in phytoplankton communities and contribute to the ecological success of harmful algal bloom (HAB) taxa. Toxin composition and content in phytoplankton are affected by a suite of environmental factors, including nutrient availability. Changes in nutrient availability can increase or decrease toxin content and alter toxin composition, depending on toxin stoichiometry and the mechanisms by which nutrient limitation affects toxin production. The studies that have assessed the effects of nutrient availability on brevetoxin content of the HAB species Karenia brevis have reported contradictory results, although there is growing support that nutrient limitation increases brevetoxin content. In this study, we assessed the effects of decreased nitrogen (N) and phosphorus (P) availability on brevetoxin content and composition of K. brevis grown in chemostats at steady state by altering the nutrient supply ratios of incoming media from the Redfield Ratio. Overall, brevetoxin content was greatest in cultures grown at the lowest rate, regardless of the nutrient supply ratio (i.e., under both Redfield and N-limiting supply ratios). Compared to cultures grown at 0.2 d⁻¹, cultures grown at 0.1 d⁻¹ exhibited 5-fold increases in intracellular toxin content. In contrast, at constant growth rates, N-limiting supply ratios decreased intracellular brevetoxin content by approximately one-third, although this result was significant only in cultures growing at the fastest rate of 0.23 d⁻¹. P-limiting supply ratios had no effect on brevetoxin content or composition. In addition, when cultures grown at rates of 0.2 d⁻¹ were supplied with balanced/Redfield N:P supply ratios, but different absolute nutrient concentrations, toxin content was greater under greater nutrient concentrations. These findings suggest that when growth rate is not nutrient limited, there is a positive relationship between nutrient availability and brevetoxin content. This work contributes to previous studies by demonstrating strong growth rates effects on brevetoxin content and that growth rate and nutrient availability can independently or together affect toxin content of K. brevis. Moreover, our work underscores the value of the chemostat as a tool to elucidate the mechanisms by which nutrient availability and growth rate affect toxin production and content of HAB species.     

OPTIMUM N:P RATIOS AND COEXISTENCE OF PLANKTONIC ALGAE1

The optimum atomic ratio of N to P, the ratio at which one nutrient limitation changes over to the other, was determined in seven species of freshwater planktonic algae. The ratio varied over a wide range among species; the average for these species was 17. If the cellular nutrient ratios in marine species are comparable with those in freshwater organisms, Redfield's ratio of 15 is remarkably close to the average. Cellular N:P ratios varied over a 24-h period under a light:dark cycle. The variation of the optimum ratio between species and diel change in cellular N:P ratios within a species could play an important role in population dynamics by enhancing the probability of coexistence of species.     

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).     

Prorocentrum minimum (Pavillard) Schiller: A review of a harmful algal bloom species of growing worldwide importance

Prorocentrum minimum (Pavillard) Schiller, a common, neritic, bloom-forming dinoflagellate, is the cause of harmful blooms in many estuarine and coastal environments. Among harmful algal bloom species, P. minimum is important for the following reasons: it is widely distributed geographically in temperate and subtropical waters; it is potentially harmful to humans via shellfish poisoning; it has detrimental effects at both the organismal and environmental levels; blooms appear to be undergoing a geographical expansion over the past several decades; and, a relationship appears to exist between blooms of this species and increasing coastal eutrophication. Although shellfish toxicity with associated human impacts has been attributed to P. minimum blooms from a variety of coastal environments (Japan; France; Norway; Netherlands; New York, USA), only clones isolated from the Mediterranean coast of France, and shellfish exposed to P. minimum blooms in this area, have been shown to contain a water soluble neurotoxic component which killed mice. Detrimental ecosystem effects associated with blooms range from fish and zoobenthic mortalities to shellfish aquaculture mortalities, attributable to both indirect biomass effects (e.g., low dissolved oxygen) and toxic effects. P. minimum blooms generally occur under conditions of high temperatures and incident irradiances and low to moderate salinities in coastal and estuarine environments often characterized as eutrophic, although they have been found under widely varying salinities and temperatures if other factors are conducive for growth. The physiological flexibility of P. minimum in response to changing environmental parameters (e.g., light, temperature, salinity) as well as its ability to utilize both inorganic and organic nitrogen, phosphorus, and carbon nutrient sources, suggest that increasing blooms of this species are a response to increasing coastal eutrophication.     

氮磷营养盐影响海水浮游硅藻种群组成的初步研究

用实验生态学方法研究了氮磷营养盐对近岸海洋硅藻组成的影响?结果表明,氮磷营养盐浓度及其比例可能对海水 游硅藻组成有着明显的影响,氮磷营养盐浓度越高,氮磷化离Ｒｅｄｆｉｅｌｄ比越远,硅藻种类越少,Ｓｈａｎｎｏｎ指数越低,这一结果在对虾养殖池浮游植物的观测中得到初步验证。     

POSITIVE FEEDBACK AND THE DEVELOPMENT AND PERSISTENCE OF ECOSYSTEM DISRUPTIVE ALGAL BLOOMS1

Harmful algal blooms (HABs) have occurred with increasing frequency in recent years with eutrophication and other anthropogenic alterations of coastal ecosystems. Many of these blooms severely alter or degrade ecosystem function, and are referred to here as ecosystem disruptive algal blooms (EDABs). These blooms are often caused by toxic or unpalatable species that decrease grazing rates by planktonic and benthic herbivores, and thereby disrupt the transfer of nutrients and energy to higher trophic levels, and decrease nutrient recycling. Many factors, such as nutrient availability and herbivore grazing have been proposed to separately influence EDAB dynamics, but interactions among these factors have rarely been considered. Here we discuss positive feedback interactions among nutrient availability, herbivore grazing, and nutrient regeneration, which have the potential to substantially influence the dynamics of EDAB events. The positive feedbacks result from a reduction of grazing rates on EDAB species caused by toxicity or unpalatability of these algae, which promotes the proliferation of the EDAB species. The decreased rates also lower grazer‐mediated recycling of nutrients and thereby decrease nutrient availability. Since many EDAB species are well‐adapted to nutrient‐stressed environments and many exhibit increased toxin production and toxicity under nutrient limitation, positive feedbacks are established which can greatly increase the rate of bloom development and the adverse effects on the ecosystem. An understanding of how these feedbacks interact with other regulating factors, such as benthic/pelagic nutrient coupling, physical forcing, and life cycles of EDAB species provides a substantial future challenge.     

Does Microorganism Stoichiometry Predict Microbial Food Web Interactions After a Phosphorus Pulse?

Knowledge of variations in microbial food web interactions resulting from atmospheric nutrient loads is crucial to improve our understanding of aquatic food web structure in pristine ecosystems. Three experiments mimicking atmospheric inputs at different nitrogen/phosphorus (N/P) ratios were performed in situ covering the seasonal biological succession of the pelagic zone in a high-mountain Spanish lake. In all experiments, abundance, biomass, algal cell biovolume, P-incorporation rates, P-cell quota, and N/P ratio of algae strongly responded to P-enrichment, whereas heterotrophic bacteria remained relatively unchanged. Ciliates were severely restricted when a strong algal exploitation of the available P (bloom growth or storage strategies) led to transient (mid-ice-free experiment) or chronic (late ice-free experiment) P-deficiencies in bacteria. In contrast, maximum development of ciliates was reached when bacteria remained P-rich (N/P < 20) and algae approached Redfield proportions (N/P approximately 16). Evidence of a higher P-incorporation rate supports the proposition that algae and bacteria shifted from a mainly commensalistic-mutualistic to a competitive relationship for the available P when bacterial P-deficiency increased, as reflected by their unbalanced N/P ratio (N/P > 20-24). Hence, the bacterial N/P ratio proved be a key factor to understand the algae-bacteria relationship and microbial food web development. This study not only demonstrates the interdependence of life history strategies, stoichiometric nutrient content, and growth but also supports the use of bacterial N/P thresholds for diagnosing ciliate development, a little-studied aspect worthy of further attention.     

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。     

Empirical models for predicting the excretion of nutrients (N and P) by aquatic metazoans: taxonomic differences in rates and element ratios

1. We developed empirical models for predicting the release of nutrients [nitrogen (N) and phosphorus (P)] by aquatic metazoans (zooplankton, mussels, benthic macroinvertebrates and fish). 2. The number of species represented in each model ranged from 9 to 74 (n = 40 – 1122), organism dry mass from 1 × 10^?5 to 8 × 10⁴ mg and water temperature from ?1.8 to 32 °C for all models. Organisms were from marine and freshwater (both lotic and lentic) environments. 3. Rates and ratios of nutrient excretion were modelled and intra‐ and intertaxon differences in excretion were examined. Rates of N and P excretion were not significantly different between marine and freshwater species within the same taxon (e.g. zooplankton). However, rates of excretion (as a function of organism dry mass and water temperature) were significantly different among different orders of zooplankton, mussels and fish. However, excretion of N was similar among different orders of benthic macroinvertebrates. 4. Detritivorous fish excreted both N and P at rates greater than all other taxa; whereas mussels excreted N and P generally at rates less than other taxa. There were no significant differences in the rate of N and P excretion between zooplankton and fish (i.e. the allometry of N and P excretion was similar between zooplankton and fish). 5. Molar N : P ratios of nutrients excreted increased with increasing organism dry mass for each group of metazoans, except for zooplankton and detritivorous fish (where N : P ratios declined with increasing organism dry mass). Molar N : P ratios in the excretions of aquatic metazoans were generally below the Redfield ratio of 16:1. 6. We examined the influence of variable abundance of zooplankton, benthic macroinvertebrates and fish on assemblage excretion rates. Rates of N and P excretion were calculated by applying our models to metazoan biomass and abundance data over seven consecutive years in two oligotrophic lakes. Rates of N and P excretion (g ha^?1 day^?1) increased linearly with increasing assemblage biomass (kg ha^?1). However, rates of N and P excretion were significantly and negatively correlated with the relative abundance of fish and positively correlated with the relative abundance of zooplankton.     

Effect of continuous nutrient enrichment on microalgae colonizing hard substrates

In order to understand the effect of changing nutrient conditions on benthic microalgae on hard substrates, in-situ experiments with artificial substrates were conducted in Kiel Fjord, Western Baltic Sea. As an extension of previous investigations, we used artificial substrates without silicate and thus were able to supply nutrient media with different Si:N ratios to porous substrates, from where they trickled out continuously. The biofilm developing on these substrates showed a significant increase in biovolume due to N + P enrichment, while Si alone had only minor effects. The stoichiometric composition of the biomass indicated nitrogen limitation during most of the year. The C:N ratios were lowered by the N + P addition. The algae were dominated by diatoms in most cases, but rhodophytes and chlorophytes also became important. The nutrient treatment affected the taxonomic composition mostly at the species level. The significance of the results with regard to coastal eutrophication is discussed.     

CULTIVATION STUDIES OF GIGARTINA SKOTTSBERGII IN SOUTHERN CHILE

Increases in population and agriculture in coastal areas can result in increased nutrient inputs and alterations in the ratios of organic to inorganic nutrients in coastal waters. Such changes in coastal nutrient regimes can affect phytoplankton community structure by creating conditions favorable for growth and dominance of algae that were not dominant before. The effect that changes in ratios and concentrations of nutrients have on toxicity of harmful algal species is not well known. There seems to be a relationship; however, between nutrient stress and toxin production among harmful phytoplankton producing low-N toxins, e.g. Diarrhetic Shellfish Poisoning (DSP) toxins. Even less is known about the relationship between organic nutrient uptake and toxin production. Benthic species and species in coastal areas are probably exposed to greater fluxes of dissolved organic nitrogen (DON). In this study, benthic and planktonic species of Prorocentrum were grown on L1 media with the sole N-source varying among treatments as nitrate, ammonium, urea, L-glutamic acid, and high molecular weight natural DON. An ELISA specific to the DSP toxins, okadaic acid and 35-methylokadaic acid, was used to determine toxin production by each species when grown on the different N sources. Preliminary results indicate that some organic forms of N support growth as well as inorganic forms for Prorocentrum minimum , P. mexicanum , and P. hoffmannianum.     

Does the Redfield ratio infer nutrient limitation in the macroalga Spirogyra fluviatilis?

1. The cellular nutrient contents of microalgae, when growing at or approaching maximum rates, approximate the Redfield C : N : P (molar) ratio of 106 : 16 : 1. Deviations from this optimal ratio can be used to infer nutrient limitation of microalgal growth. However, this ratio may not be applicable to macroalgae, which are distinguished from microalgae by forming a thallus that is a discrete structure visible to the naked eye. The utility of the Redfield ratio to infer nutrient limitation of the growth of macroalgae was tested for Spirogyra fluviatilis in a field experiment conducted in tropical Australia. 2. The optimal cellular C : N : P ratio for S. fluvialitis was estimated by means of in situ nutrient addition. This was compared with S. fluvialitis cellular ratios determined from eight sites with a wide range of soluble N concentrations (<1–90 μg L^?1), a smaller range of soluble P concentrations (5–12 μg L^?1), and soluble molar N : P ratios of 0.11– 27. 3. Spirogyra fluviatilis had an optimal molar C : N : P ratio of 1800 : 87 : 1 which differs substantially from the Redfield ratio, and suggests that the latter ratio is not applicable to this macroalga. Concentrations of N and P in the river deviated from the optimal N : P ratio of 87 : 1, inferring nutrient limitation of growth. 4. C : P and C : N ratios of S. fluviatilis varied in accordance with general stoichiometric relationships for autotrophs under nutrient limitation of growth. Ratios of C : P and C : N increased, respectively, with increased severity of P‐ and N‐limitation. Additionally, C : P ratios increased with increased N : P ratios, whilst the C : N ratio increased with decreased N : P ratios. The C : N molar ratio however was an insensitive indicator of nutrient depletion compared with the C : P ratio. Under N‐limitation of growth, luxury amounts of P were stored by S. fluviatilis. 5. In aquatic environments where macroalgae are sufficiently abundant to be sampled, their cellular carbon, nitrogen and phosphorus stoichiometry can be used to infer nutrient limitation of growth when their optimal C : N : P ratio is known.     

Nitrogen, phosphorus and potassium nutritional status of semiarid steppe grassland in Inner Mongolia

In grazed semiarid steppe ecosystems, much attention has been paid to aspects of growth limitation by water. So far, potential limitation of primary production by plant nutrients was rarely considered. This knowledge is essential for identification of sustainable land-use practices in these large and important ecosystems on the background of over-exploitation and climate change. In the present study plant nutrient concentrations and ratios were investigated with factorial additions of water and N fertilizer at two sites with contrasting soil nutrient availability. Combined analysis of nutrient concentrations, contents, biomass production, and plant N:P ratios consistently confirmed primary growth limitation by water and a strong N limitation when sufficient amounts of water were supplied. P limitation only occurred at the site with low P availability when in addition to the natural supply, water and N fertilizer were given. According to reported thresholds of N:K and K:P ratios, K was not limiting in any plot. The observed nutritional patterns in the plant community were related to the dynamics of species composition and their specific nutrient status. Stipa grandis had the highest N:P ratio whereas Artemisia frigida showed lowest N:P. These nutrient characteristics were related to growth strategies of dominant species. Accordingly, the relative biomass contribution of S. grandis and A. frigida strongly affected the nutrient status of the plant community. Plant N:P ratios indicate the relative limitation by N or P in the semiarid grasslands under sufficient water supply, but other methods of nutritional diagnosis should be used when plant N:P ratios remain below critical values.     

Grazer species effects on epilithon nutrient composition

1. Field and laboratory experiments were conducted to investigate the excretion stoichiometry of nitrogen (N) and phosphorus (P) of two benthic macroinvertebrate grazers, the crayfish Orconectes propinquus and the snail Elimia livescens, that differ in body stoichiometry (mean body molar N : P 18 and 28, respectively). Crayfish excretion had a significantly higher ammonium : soluble reactive phosphorus (SRP) ratio in the laboratory and in three natural streams than did snails, as predicted by ecological stoichiometry theory. 2. In greenhouse recirculating artificial streams, treatments consisting of crayfish, snails, or no grazers were used to examine responses in dissolved nutrient concentrations and epilithon nutrient composition and limitation. SRP concentrations depended upon the grazer species, with the snail treatment having a higher SRP concentration than other treatments (P < 0.05). Dissolved inorganic N was not affected by grazers, but appeared to be rapidly incorporated in epilithon. 3. Epilithon N content was dependent upon the grazer species present, with the crayfish treatment having a significantly higher N content than other treatments (P = 0.001). No grazer species effects on epilithon P content were found. However, both grazer treatments had significantly lower epilithon P content than the no‐grazer treatment. 4. Traditionally, studies have focused on how grazer‐induced structural changes to epilithon can alter epilithon nutrient dynamics, but this structural mechanism could not solely explain differences in epilithon nutrient contents and ratios in the present study. Our results rather suggest that benthic grazers can alter epilithon nutrient composition and limitation via nutrient excretion. Consequently, macroinvertebrate grazers may serve as ‘nutrient pumps’ that partly regulate the availability of nutrients to algae in stream ecosystems.     

Nitrogenous Nutrients Promote the Growth and Toxicity of Dinophysis acuminata during Estuarine Bloom Events

Diarrhetic Shellfish Poisoning (DSP) is a globally significant human health syndrome most commonly caused by dinoflagellates within the genus Dinophysis. While blooms of harmful algae have frequently been linked to excessive nutrient loading, Dinophysis is a mixotrophic alga whose growth is typically associated with prey availability. Consequently, field studies of Dinophysis and nutrients have been rare. Here, the temporal dynamics of Dinophysis acuminata blooms, DSP toxins, and nutrients (nitrate, ammonium, phosphate, silicate, organic compounds) were examined over four years within two New York estuaries (Meetinghouse Creek and Northport Bay). Further, changes in the abundance and toxicity of D. acuminata were assessed during a series of nutrient amendment experiments performed over a three year period. During the study, Dinophysis acuminata blooms exceeding one million cells L-1 were observed in both estuaries. Highly significant (p<0.001) forward stepwise multivariate regression models of ecosystem observations demonstrated that D. acuminata abundances were positively dependent on multiple environmental parameters including ammonium (p = 0.007) while cellular toxin content was positively dependent on ammonium (p = 0.002) but negatively dependent on nitrate (p<0.001). Nitrogen- (N) and phosphorus- (P) containing inorganic and organic nutrients significantly enhanced D. acuminata densities in nearly all (13 of 14) experiments performed. Ammonium significantly increased cell densities in 10 of 11 experiments, while glutamine significantly enhanced cellular DSP content in 4 of 5 experiments examining this compound. Nutrients may have directly or indirectly enhanced D. acuminata abundances as densities of this mixotroph during experiments were significantly correlated with multiple members of the planktonic community (phytoflagellates and Mesodinium). Collectively, this study demonstrates that nutrient loading and more specifically N-loading promotes the growth and toxicity of D. acuminata populations in coastal zones.     

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l⁻¹ and between 400 and 1000 μg TN l⁻¹.