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Summary: The phytoplankton Phaeocystis globosa plays an important role in sulfur cycling and climate control, and can develop harmful algal blooms (HABs). Here we report a chromosome-scale reference genome assembly of P. globosa, which enable in-depth analysis of molecular underpinnings of important ecological characteristics. Comparative genomic analyses detected two-rounds of genome duplications that may have fueled evolutionary innovations. The genome duplication may have resulted in the formation of dual HiDP and LoDP dimethylsulphoniopropionate (DMSP) biosynthesis pathways in P. globosa. Selective gene family expansions may have strengthened biological pathways critical for colonial formation that is often associated with the development of algal blooms. The copy numbers of rhodopsin genes are variable in different strains, suggesting that rhodopsin genes may play a role in strain-specific adaptation to ecological factors. The successful reconstruction of the P. globosa genome sets up an excellent platform that facilitates in-depth research on bloom development and DMSP metabolism.

Phaeocystis globosa is an important harmful algal species that is globally distributed. Previous studies have indicated that P. globosa preferentially uptake nitrate, but the underlying mechanism is still unclear. To further verify this preference and reveal the underlying mechanism, the assimilation rates of nitrate and ammonium by P. globosa at different concentrations was quantitatively studied by using a nitrogen stable isotope (15N) tracer technique, and the regulatory mechanism was determined from the physiological and biochemical responses. The findings revealed that the preferential assimilation of nitrate by P. globosa was influenced by the ambient ammonium concentration. When the ambient concentration of ammonium was less than approximately 3.5 μmol·L−1, the assimilation rates of nitrate form P. globosa were as high as 1.05 × 10−5 μmol·L−1·h−1·cell−1. Even though the nitrate assimilation in P. globosa was inhibited at ammonium concentrations greater than this threshold, nitrate assimilation could not be completely suppressed. The activity of NR and the expression of related genes in P. globosa were also affected by ammonium. In addition, 15N signals originally labeled nitrate accumulated in ammonium. This indicated that P. globosa was likely to reduce excess nitrate to ammonium and subsequently release it into the substrate, which might be an important energy dissipation mechanism for P. globosa. These results expand the classical understanding of the utilization of nitrogen nutrients by marine phytoplankton, and offer valuable resources for comprehending the mechanism of harmful algal blooms.

Modified clay (MC) is an effective and eco-friendly material that is widely used for controlling harmful algae blooms (HABs). In this study, a new method—single source with series modifications (SSSM)—was optimized for preparing a high-quality MC, and aluminum-rich clay (kaolin) was activated with a multistep treatment that included calcination (Cal), acid etching with alkali neutralization (Al–An), and raw clay supplementation and aging (RSA). The results showed that the removal efficiency of Prorocentrum donghaiense by the SSSM–MC exceeded 85%; Cal, Al–An, and RSA were indispensable processing steps for preparing the best MC with the SSSM method. The changes occurring in the mineral structure and chemical composition of kaolin treated by the different steps were characterized by FT-IR, XRD, 27Al-NMR and Al-Ferron. Surface dehydroxylation and disruption of the kaolin lattice after calcination significantly changed the aluminum structure from AlVI to AlV and AlIV. The relatively reactive AlV was readily released from the kaolin lattice with acid etching, and it subsequently reacted with external OH– to generate different polyhydroxyaluminum species because alkali was added during the Al–An step. The acidity of the above complex was further decreased by RSA, greatly increasing the yield of SSSM–MC while assuring the quality.

Eutrophication in aquaculture areas concurrently leads to a high incidence of dissolved oxygen deficiency and toxic algal blooms. The combined effects of hypoxia and typical toxic algae on cultured organisms should be given sufficient consideration. Abalone breeding in China has greatly suffered from hypoxia and toxic Karenia mikimotoi blooms for many years. In this study, the individual and combined effects of the toxic dinoflagellate, K. mikimotoi, and hypoxia on juvenile abalone were determined based on abalone survival and oxidative stress indicators in their gills, hepatopancreas and hemolymph. The results showed that at a density of 106 to 3×107 cells/L, K. mikimotoi alone had a negligible influence on the survival of juvenile abalone under sufficient dissolved oxygen (DO) conditions. The 24 h-half lethal concentration (LC50) of DO alone for juvenile abalone was 0.75 mg/L in seawater. When K. mikimotoi was added at a density of 3×106 cells/L, the 24 h-LC50 of DO for juvenile abalone significantly increased to 2.59 mg/L, indicating obvious synergistic effects. The individual effects of hypoxia or K. mikimotoi on the oxidative stress indicators were limited, and only the superoxide dismutase (SOD) activity in the abalone gills significantly decreased under K. mikimotoi stress. However, the combined stress of hypoxia and K. mikimotoi led to significant changes in the antioxidant indicators in all tested tissues. The SOD activity in gills and hepatopancreas decreased, while the SOD and catalase (CAT) activity and malondialdehyde (MDA) content in the hemolymph increased due to the combined stress of hypoxia and K. mikimotoi. These results illustrated that the synergistic effects of hypoxia and K. mikimotoi caused serious oxidative damage in abalone and that the hemolymph exhibited greater sensitivity than did the gills and hepatopancreas. Further investigation found that K. mikimotoi increased the oxygen consumption rate in abalone and that hypoxia enhanced the hemolytic toxicity of K. mikimotoi. These results revealed that hypoxia and typical toxic algae cause synergistic harm to cultured organisms, which is expected to provide a new understanding of the destructive mechanisms of typical toxic algal blooms in aquacultural areas.

Abstract Harmful algal blooms (HABs) have occurred more frequently in recent years. In this study, to investigate their potential impact in the Beibu Gulf, short‐read and long‐read metabarcoding analyses were combined for annual marine phytoplankton community and HAB species identification. Short‐read metabarcoding showed a high level of phytoplankton biodiversity in this area, with Dinophyceae dominating, especially Gymnodiniales. Multiple small phytoplankton, including Prymnesiophyceae and Prasinophyceae, were also identified, which complements the previous lack of identifying small phytoplankton and those unstable after fixation. Of the top 20 phytoplankton genera identified, 15 were HAB‐forming genera, which accounted for 47.3%–71.5% of the relative abundance of phytoplankton. Based on long‐read metabarcoding, a total of 147 OTUs (PID > 97%) belonging to phytoplankton were identified at the species level, including 118 species. Among them, 37 species belonged to HAB‐forming species, and 98 species were reported for the first time in the Beibu Gulf. Contrasting the two metabarcoding approaches at the class level, they both showed a predominance of Dinophyceae, and both included high abundances of Bacillariophyceae, Prasinophyceae, and Prymnesiophyceae, but the relative contents of the classes varied. Notably, the results of the two metabarcoding approaches were quite different below the genus level. The high abundance and diversity of HAB species were probably due to their special life history and multiple nutritional modes. Annual HAB species variation revealed in this study provided a basis for evaluating their potential impact on aquaculture and even nuclear power plant safety in the Beibu Gulf.

Herein, the algicidal effects of Brevibacillus laterosporus on typical red tide organisms were investigated. Through comparative analysis of the fermentation solution, sterile filtrate and bacterial body, it was found that this strain mainly exerts its algicidal function by secreting algicidal substances. In this paper, we established a method for extracting the algicidal substances of B. laterosporus and systematically investigated their features and effect on Heterosigma akashiwo. The results showed that the algicidal substances are a mixture of compounds all with molecular weights less than 3500 Da, with those below 100 Da producing 45% of the algicidal effect; these are more polar and best extracted with methanol. The algicidal substances are stable over a range of temperatures (-80.0 to 70.0°C) and pH values (4.0-8.0). The algicidal substances caused most H. akashiwo cells to rupture within 24 h. In the remaining cells, the algicidal substances activated the antioxidant system and reduced their metabolic activity, leading to apoptosis, as observed by cell crumbling and a reduction in membrane potential. The responses of different algal cell surface structures to the algicidal substances were also compared and analysed. It was concluded that these algicidal substances can act on the cell membrane and change its permeability, allowing entry of the algicidal substances to produce an algicidal effect.

Paralytic shellfish toxins (PSTs), produced by Alexandrium pacificum in the marine environment, are a group of potent neurotoxins which specifically block voltage-gated sodium channels in excitable cells. During the toxigenic A. pacificum blooms outbreaks, PSTs can be accumulated through the food chain and finally enter the human body, posing a significant threat to human health and safety. This study experimented with a novel type of oxidized modified clay, potassium peroxymonosulfate modified clay (PMPS-MC), which could remove A. pacificum cells as well as reduce intracellular and extracellular PSTs toxicity rapidly. For the extracellular PSTs, its content decreased to below the detection limit rapidly through oxidative degradation within 15 min of 10 mg/L PMPS-MC treatment. Whereafter, although the residual cells in water column and some viable cells in flocculated sediment continued to secrete toxins, the extracellular PSTs content and toxicity in the PMPS-MC treatment groups remained significantly lower than those in the control group. For the intracellular PSTs, PMPS-MC might induce the transformation of more toxic GTX1&4 to less toxic GTX2&3 and C1&2, resulting in intracellular PSTs toxicity reduced within 15 min. In addition, intracellular PSTs content and toxicity in the PMPS-MC treatment groups were consistently lower than the control group within 48 h, possibly by inhibiting the A. pacificum cells growth. These results will provide a scientific basis for the field application of modified clay to control A. pacificum blooms.

Red tide is a kind of marine disaster caused by the accumulation or proliferation of microalgae and other organisms in a short period of time, and utilizing modified clay to control and inhibit red tide is the preferred method. Among them, the application potential of organic-modified clay is high; unlike inorganic and microbial modifications, it has a broad-spectrum removal capacity on red tide algae at extremely low dosages. However, it has some disadvantages such as severe toxicity and high residual turbidity, leading to several limitations in its practical application. Therefore, it has become urgent to select organic-modified reagents with higher efficiency, weaker toxicity and lower residual turbidity. In this study, the typical red tide alga——Prorocentrum donghaiense was selected to detect the removal capacity of Polydimethyl diallyl ammonium chloride (PDMDAAC) modified clay (MP) by comparing with the Hexadecyl trimethyl ammonium bromide (HDTMA) modified clay (MH). Not only the physiological stress and flocculation effects of two modified clays on microalgae had been discussed, but also the properties of the modified clays had been characterized in this study. The results showed that the low degree of oxidative stress and less damage to the cell membrane make MP more environmentally friendly, PDMDAAC can remove microalgae at a low dose (2 mg/L) and quickly clarify the water by significantly enhancing the flocculation capacity of clay. In addition to discussing the removal mechanism of two modified clays on microalgae, schematic diagrams of the pathways were drafted. This study will provide support for the development of organic-modified clay.

Nutrients and phytoplankton associated with mariculture development are important concerns globally, as they can significantly impact water quality and aquaculture yield. Currently, there is still insufficient information regarding the variations in nutrients and phytoplankton community of intensive mariculture systems, and effective treatment is lacking. Here, based on consecutive daily monitoring of two Litopenaeus vannamei ponds from July to October, the dynamic variations in nutrients and phytoplankton were elucidated. In addition, modified clay (MC) method was adopted to regulate the nutrients and phytoplankton community. The temporal variations in organic and inorganic nutrients presented fluctuating upward trends. Notably, organic nutrients were the dominant species, with average proportions of TON/P in TN/P were as high as 75.29% and 87.36%, respectively. Furthermore, a marked increase in the ratios of dinoflagellates to diatoms abundance were also observed in the control pond, concurrently with dominant organic nutrients, ascending N/P ratio and decreasing Si/N and Si/P ratios. In the MC-regulated pond, MC reduced the contents of both organic and inorganic nutrients. Furthermore, a distinct change pattern of dominant phytoplankton community occurred, with green algae becoming the most abundant phytoplankton in the MC-regulated pond. This study can provide new insights into an effective treatment for managing water quality and maintaining sustainable mariculture development.

Antibiotics have been identified as emerging pollutants, and they increasingly threaten the safety of mariculture; thus, effective treatment methods are urgently needed. In this study, Fe(II)-peroxymonosulfate-modified clay (Fe-PMS-MC), an effective mineral complex for harmful algal bloom control, was tested for sulfamethoxazole (SMX) removal from seawater. The results showed that SMX removal efficiency increased gradually from 42.3% to 100% in seawater in 6 h as the Fe-PMS-MC dose increased from 0.1 g/L to 1.0 g/L; this removal was more efficient than that in freshwater. Both an increase in temperature and the addition of inorganic anions such as Cl−, HCO3−, and SO42− improved the SMX removal efficiency; in contrast, there was no noticeable influence of a pH change from 3.0 to 9.0 or of Heterosigma akashiwo biomass addition from 103 cells/mL to 105 cells/mL. Oxidation was the main mechanism for the removal of SMX by Fe-PMS-MC treatment, and the mechanisms included radical oxidation and non-radical oxidation. PMS was the key component in Fe-PMS-MC for the production of radicals, which increased with temperature. Similarly, both radical oxidation and non-radical oxidation increased when PMS was attacked by Cl−, HCO3−, and SO42− in seawater.

In this paper, atomic force microscopy was employed to study the harmful algal bloom species Prorocentrum donghaiense for the first time. Cells were immobilized in pores of polycarbonate membrane to keep moist and to acquire images of P. donghaiense at different scales. Typical ultrastructures, such as knob-like spines and valve pores, were observed on cell surfaces. These structures had similar characteristics to those observed on scanning electron microscopy images. Moreover, the height and spacing of typical nanostructure, and nanomechanical parameters such as adhesion and elasticity, were also quantified by AFM. Additionally, the changes in cell surface nanostructure and nanomechanical characteristics under nitrogen limitation were further studied. Compared with the cells under normal culture conditions, the cell surface roughness and adhesion decreased, and the elastic modulus increased for cells under nitrogen limitation. Potential changes in the ability of P. donghaiense cells to perform normal physiological functions are reflected by changes in cell surface parameters, including cell surface roughness, cell surface adhesion, and cell surface elasticity. The results of this study reveal how P. donghaiense responds to changes in the external environment under approximately physiological conditions from the perspective of changes in cell surface nanostructures and nanomechanical characteristics and provide a new understanding of its cell biology.

Modified clay (MC) technology is the most effective method to control harmful algal blooms (HABs) and has been widely testified in many countries in recent decades. Although dissolved algal organic matter (dAOM) has been found to be abundant in HAB-affected waters, little is known about its effect on MC flocculation. Prorocentrum donghaiense was chosen as the model organism in this study. The flocculation of MC with different concentrations of dAOM was observed by particle image velocimeter, and the removal efficiency of MC to microalgae was determined using a fluorometer. The results showed that a small amount of dAOM resulted in faster flocculation, larger flocs, higher floc strength and better floc regeneration capacity, and the removal efficiency of microalgae by MC could reach about 80%. However, large amounts of dAOM produced during the occurrence of HABs could inhibit the flocculation of MC, and the removal efficiency of microalgae was only about 35%. Furthermore, with the increase of dAOM, the zeta potential of MC particles decreases from 1.56 mV to −18.9 mV, and the repulsive force between the particles also increases. The examination of 18-angle laser light scattering gel permeation chromatography and specific ultraviolet absorption (SUVA254) revealed that some hydrophobic organic macromolecules preferred to attach to MC and increase MC flocculation at a tiny amount of dAOM, while a higher concentration of dAOM would inhibits flocculation by reducing surface activity and intergranular repulsion. According to the findings of this study, the dosage of MC must be increased in order to obtain the best algal removal efficiency with MC.

Algicidal bacteria are important in the control of toxic dinoflagellate blooms, but studies on the environmental behavior of related algal toxins are still lacking. In this study, Bacillus subtilis S3 (S3) showed the highest algicidal activity against Alexandrium pacificum (Group IV) out of six Bacillus strains. When treated with 0.5% (v/v) S3 bacterial culture and sterile supernatant, the algicidal rates were 69.74% and 70.22% at 12 h, respectively, and algicidal substances secreted by S3 were considered the mechanism of algicidal effect. During the algicidal process, the rapid proliferation of Alteromonas sp. in the phycosphere of A. pacificum may have accelerated the algal death. Moreover, the algicidal development of S3 released large amounts of intracellular paralytic shellfish toxins (PSTs) into the water, as the extracellular PSTs increased by 187.88% and 231.47% at 12 h, compared with the treatment of bacterial culture and sterile supernatant at 0 h, respectively. Although the total amount of PSTs increased slightly, the total toxicity of the algal sample decreased as GTX1/4 was transformed by S3 into GTX2/3 and GTX5. These results more comprehensively reveal the complex relationship between algicidal bacteria and microalgae, providing a potential source of biological control for harmful algal blooms and toxins.

Blooms of the toxic dinoflagellate Karenia mikimotoi could threaten the survival of marine life, and modified clay (MC) is considered a promising method for the control of harmful algal blooms. Here, using marine medaka as the model organism, the toxicity of K. mikimotoi before and after MC disposal was investigated. The results showed that only a certain density of intact K. mikimotoi cells could cause obvious damage to fish gills and lead to rapid death. A systematic analysis of morphology, physiology, and molecular biology parameters revealed that the fish gills exhibited structural damage, oxidative damage, osmotic regulation impairment, immune response activation, and signal transduction enhancement. MC can flocculate K. mikimotoi rapidly in water and reduce its toxicity by reducing the density of intact algae cells and hemolytic toxicity. The results indicate that MC is an effective and safe method for controlling K. mikimotoi blooms.

An outbreak of harmful algal blooms (HABs) often leads to the death of fish and other marine organisms and causes serious losses to human economic activities. Modified clay (MC) technology is an effective way to control HABs. Although the material and preparation process are based on the premise of green and safety, the potential eco-environmental impacts of MC on non-target organisms should still have to receive attention before field applications can occur. Therefore, the effects of one commonly used modified clay, polyaluminum chloride-modified clay (PAC-MC), on the survival, growth, and oxidative stress of the marine medaka (Oryzias melastigma) were studied. The toxicity test results showed that the 96-h median lethal concentration (96-h LC50) of PAC-MC for newly hatched medaka larvae was 5.204 g/L, which was much higher than the concentration used on site (4–10 t/km2). Within the concentration range of PAC-MC used in this experiment (≤2 g/L), the morphology, heart rate, growth, and aluminum content of larvae did not change with the increase in the modified clay concentration. Low concentrations of PAC-MC (≤0.5 g/L) did not significantly affect catalase (CAT) activity, superoxide dismutase (SOD) activity, peroxidase (POD) activity, and the content of malondialdehyde (MDA), but higher concentrations of PAC-MC (such as 2 g/L) caused oxidative damage to the larvae and increased the antioxidant enzyme activity of the larvae. The present study revealed that under an effective dosage for treating harmful algal blooms on site, PAC-MC had no adverse effects on the survival, growth, oxidative stress, and aluminum content of the newly hatched marine medaka, which provides a scientific basis for the field application of modified clay.

In this study, Karenia brevis 165 (K. brevis 165), a Chinese strain, was used to research brevetoxin (BTX) metabolites. The sample pretreatment method for the enrichment of BTX metabolites in an algal culture medium was improved here. The method for screening and identifying intracellular and extracellular BTX metabolites was established based on liquid chromatography-time-of-flight mass spectrometry (LC-ToF-MS) and liquid chromatography triple quadrupole tandem mass spectrometry (LC-QqQ-MS/MS). The results show that the recovery rates for BTX toxins enriched by a hydrophilic–lipophilic balance (HLB) extraction column were higher than those with a C18 extraction column. This method was used to analyze the profiles of extracellular and intracellular BTX metabolites at different growth stages of K. brevis 165. This is the first time a Chinese strain of K. brevis has been reported that can produce toxic BTX metabolites. Five and eight kinds of BTX toxin metabolites were detected in the cell and culture media of K. brevis 165, respectively. Brevenal, a toxic BTX metabolite antagonist, was found for the first time in the culture media. The toxic BTX metabolites and brevenal in the K. brevis 165 cell and culture media were found to be fully proven in terms of the necessity of establishing a method for screening and identifying toxic BTX metabolites. The results found by qualitatively and quantitatively analyzing BTX metabolites produced by K. brevis 165 at different growth stages show that the total toxic BTX metabolite contents in single cells ranged between 6.78 and 21.53 pg/cell, and the total toxin concentration in culture media ranged between 10.27 and 449.11 μg/L. There were significant differences in the types and contents of toxic BTX metabolites with varying growth stages. Therefore, when harmful algal blooms occur, the accurate determination of BTX metabolite types and concentrations will be helpful to assess the ecological disaster risk in order to avoid hazards and provide appropriate disaster warnings.

Phosphine, which is released continuously from sediment, can affect the eco-physiological strategies and molecular responses of phytoplankton. To examine the effects of phosphine on phosphorus uptake and utilization in Thalassiosira pseudonana, we examined the transcriptional level of the phosphate transporter gene (TpPHO) and the activity of alkaline phosphatase (AKP) in relation to supplement of various concentrations of phosphine. TpPHO expression was markedly promoted by phosphine in both the phosphate-deficient and phosphate-4 µM culture. However, high phosphine concentrations can inhibit TpPHO transcription in the declining growth phase. AKP activity was also higher in the phosphine treatment groups than that of the control. It increased with increasing phosphine concentration in the range of 0 to 0.056 µM but was inhibited by higher levels of phosphine. These responses revealed that phosphine can affect phosphate uptake and utilization in T. pseudonana. This result was consistent with the effect of phosphine on algal growth, while TpPHO expression and AKP were even more sensitive to phosphine than algal growth. This work provides a basic understanding for further research about how phosphine affects phytoplankton.

Hypoxia has become an overwhelming phenomenon in the coastal environment off the Changjiang (Yangtze River) estuary. Seasonal data were obtained in the Changjiang estuary during 2003, 2004, 2005, and 2009 and during an additional cruise between April and May in 2007 on the dissolved oxygen distribution of the bottom water, on hypoxic events, and on their potential causes. Dissolved oxygen was less than 3.0 mg L-1 in the summer and less than 5.0 mg L-1 in the spring and autumn in the topographic trough and was generally lower than that in the surrounding bottom water of the Changjiang estuary. In the summer, there was a positive correlation (r2 = 0.57) between dissolved oxygen concentration and temperature, contrary to the observations in the autumn and winter. A strong halocline and thermocline appeared in the spring and summer. A change in temperature structure was observed in an additional cruise in 2007; the vertical transport of dissolved oxygen flux, which was restricted by stratification, reached 2.77 g m-2 d-1 Large nutrient inputs and phytoplankton seem to contribute to hypoxia in the Changjiang estuary. [ABSTRACT FROM AUTHOR]

Abstract: Fish Lateolabrax japonicus were exposed to 0.1 and 1mg/L of anion surfactant sodium dodecylbenzene sulfonate (SDBS) and to 2 and 20μg/L of benzo[a]pyrene (B[a]P) for 6, 12, and 18 days, with control and solvent control groups. Liver antioxidant enzymes, including superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPx), reduced glutathione (GSH), and glutathione S-transferase (GST), were determined; brain acetylcholinesterase (AChE) and liver inducible nitric oxide synthase (iNOS) activities were also measured. The results indicated that (1) L. japonicus avoided oxidative damage through antioxidant systems; (2) SOD, GPx, and GSH were induced, and GST was inhibited and then induced by B[a]P exposure; and (3) CAT, GPx, and AChE were induced while iNOS was inhibited, and GST was induced and then inhibited by SDBS stress in experimental period. [Copyright &y& Elsevier]

Eutrophication has become increasingly serious and noxious algal blooms have been of more frequent occurrence in the Yangtze River Estuary and in the adjacent East China Sea. In 2003 and 2004, four cruises were undertaken in three zones in the estuary and in the adjacent sea to investigate nitrate (NO3–N), ammonium (NH4–N), nitrite (NO2–N), soluble reactive phosphorus (SRP), dissolved reactive silica (DRSi), dissolved oxygen (DO), phytoplankton chlorophyll a (Chl a) and suspended particulate matter (SPM). The highest concentrations of DIN (NO3–N+NH4–N+NO2–N), SRP and DRSi were 131.6, 1.2 and 155.6 μM, respectively. The maximum Chl a concentration was 19.5 mg m−3 in spring. An analysis of historical and recent data revealed that in the last 40 years, nitrate and SRP concentrations increased from 11 to 97 μM and from 0.4 to 0.95 μM, respectively. From 1963 to 2004, N:P ratios also increased from 30–40 up to 150. In parallel with the N and P enrichment, a significant increase of Chl a was detected, Chl a maximum being 20 mg m−3, nearly four times higher than in the 1980s. In 2004, the mean DO concentration in bottom waters was 4.35 mg l−1, much lower than in the 1980s. In comparison with other estuaries, the Yangtze River Estuary was characterized by high DIN and DRSi concentrations, with low SRP concentrations. Despite the higher nutrient concentrations, Chl a concentrations were lower in the inner estuary (Zones 1 and 2) than in the adjacent sea (Zone 3). Based on nutrient availability, SPM and hydrodynamics, we assumed that in Zones 1 and 2 phytoplankton growth was suppressed by high turbidity, large tidal amplitude and short residence time. Furthermore, in Zone 3 water stratification was also an important factor that resulted in a greater phytoplankton biomass and lower DO concentrations. Due to hydrodynamics and turbidity, the open sea was unexpectedly more sensitive to nutrient enrichment and related eutrophication processes. [ABSTRACT FROM AUTHOR]