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1.
The parasitic dinoflagellates in the genus of Amoebophrya can infect broad ranges of planktonic dinoflagellates, and transform algal biomass into organic matter that can be recycled within the planktonic community. The ecological significance of Amoebophrya spp. during harmful algal bloom (HAB) events was gradually recognized along with revelation of its host specificity and diversity in picoplankton communities. The eutrophicated coastal waters of China are frequently affected by HABs, particularly in Changjiang (Yangtze River) estuary and the adjacent East China Sea; while, no research has been conducted to explore the ecological roles of parasitism during HAB events and the related dinoflagellate bloom dynamics. For the first time, we confirmed the presence of Amoebophrya infections in the planktonic community of this region; six species of dinoflagellates were infected, including Ceratium tripos, Scrippsiella trochoidea, Gonyaulax spinifera, Gymnodinium sp., Gonyaulax sp. and an Alexandrium sp. Molecular sequences retrieved from environmental water samples revealed high genetic diversity of Amoebophryidae-like organisms in the water column. Amoebophrya-infected dinoflagellates were only observed in high salinity (>20) stations suggesting that salinity may be a factor limiting the distribution of Amoebophyra infections in natural environment. Whereas, no evidence of Amoebophrya infection was observed in the bloom-forming species Karenia mikimotoi, suggesting that K. mikimotoi in this region was likely free of Amoebophridae infection.  相似文献   

2.
The endoparasitic dinoflagellate Amoebophrya infects a number of free‐living marine dinoflagellates, including harmful algal bloom species. The parasitoid eventually kills its host and has been proposed to be a significant loss factor for dinoflagellate blooms in restricted coastal waters. For several decades, the difficulties of culturing host‐parasitoid systems have been a great obstacle for further research on the biology of Amoebophrya. Here, we established an Akashiwo sanguineaAmoebophrya sp. coculture from Chinese coastal waters and studied the parasitoid's generation time, dinospore survival and infectivity, as well as its host specificity. The lifespan of Amoebophrya sp. ex. A. sanguinea was approximately 58 h. The infective dinospores can survive up to 78 h in ambient waters but gradually lose their infectivity. The parasitoid was unable to infect other dinoflagellate species, its infection rate reached as high as 91% when the ratio of dinospores to host cells was 20:1. The high infectivity of dinospores suggests that the Amoebophrya strain was capable of removing a considerable fraction of host biomass within a short period, but that it is probably unable to maintain high infection levels under nonbloom conditions of its host, due to limited survival and time constraints in encountering host cells.  相似文献   

3.
Amoebophrya is a marine parasite recently found to infect and kill bloom-forming dinoflagellates in the California Current System (CCS). However, it is unknown whether parasitism by Amoebophrya can control dinoflagellate blooms in major eastern boundary upwelling systems, such as the CCS. We quantified the abundance of a common bloom-forming species Akashiwo sanguinea and prevalence of its parasite (i.e., % infected cells) in surface water samples collected weekly from August 2005 to December 2008 at the Santa Cruz Wharf (SCW), Monterey Bay, CA. Additionally, we measured physical and chemical properties at the SCW and examined regional patterns of wind forcing and sea surface temperature. Relative abundance of the net phytoplankton species was also analyzed to discern whether or not parasitism influences net phytoplankton community composition. Epidemic infection outbreaks (>20% parasite prevalence in the host species) may have contributed to the end or prevented the occurrence of A. sanguinea blooms, whereas low parasite prevalence was associated with short-term (≤2 weeks) A. sanguinea blooms. The complete absence of parasitism in 2007 was associated with an extreme A. sanguinea bloom. Anomalously strong upwelling conditions were detected in 2007, suggesting that A. sanguinea was able to outgrow Amoebophrya and ‘escape’ parasitism. We conclude that parasitism can strongly influence dinoflagellate bloom dynamics in upwelling systems. Moreover, Amoebophrya may indirectly influence net phytoplankton species composition, as species that dominated the net phytoplankton and developed algal blooms never appeared to be infected.  相似文献   

4.
The dinoflagellate Akashiwo sanguinea is a well known, cosmopolitan harmful microalga that frequently forms harmful algal blooms (HABs) in marine estuaries from temperate to tropical waters, and has posed a severe threat to fish, shellfish, and sea birds. Therefore, it is important to understand the ecology of this species, particularly the mechanisms regulating its ubiquitous geographic distribution and frequent recurrence of. To date, the mechanisms regulating distribution and recurrence of HABs by this species have been poorly understood. While resting cyst production can play a central role in the geographic expansion and initiation of HABs, studies of the life cycle of this alga, including cyst production, have been lacking. Here, we demonstrate that A. sanguinea produces sexual resting cysts homothallically. We present evidence for cell pairs in sexual mating, biflagellated planozygote formation, and cysts of different morphologies, and we describe time series for germination of cysts to germlings with two longitudinal flagella, along with studies of possible factors affecting cyst production. Phylogenetic analysis of large sub‐unit rDNA sequences revealed a monophyly of this species and thus possibly a recent common ancestor for all global populations. The discovery of resting cyst production by A. sanguinea suggests its frequent recurrence of blooms and global distribution may have been facilitated by the natural and anthropogenic transport of resting cysts.  相似文献   

5.
Several algal species responsible for harmful algal blooms (HABs), such as Alexandrium fundyense, are mixotrophic under certain environmental conditions. The ability to switch between photosynthetic and heterotrophic modes of growth may play a role in the development of HABs in coastal regions. We examined the influence of humic dissolved organic matter (HDOM) derived from terrestrial (plant/soil) and microbial sources on the growth of A. fundyense. We found that a terrestrially derived HDOM, Suwannee River humic acid (SRHA), did enhance A. fundyense growth; however, a microbially derived HDOM, Pony Lake fulvic acid (PLFA) did not enhance growth. A. fundyense grows in association with bacteria in culture and we observed that bacterial cell densities were much lower in A. fundyense cultures than in bacteria‐only cultures, consistent with bacterial grazing by A. fundyense in culture. In bacteria‐only cultures with added algal exudates (EX), the addition of PLFA and SRHA resulted in a slight increase in bacterial cell density compared to cultures without HDOM added. Changes over time in the chemical quality of the HDOM in the A. fundyense cultures reflected contributions of microbially derived material with similar characteristics as the PLFA. Overall, these results suggest that the chemical differences between SRHA and PLFA are responsible for the greater effect of SRHA on A. fundyense growth, and that the differential effect is not a result of an effect on the growth of associated bacteria.  相似文献   

6.
Amoebophrya is an obligate endoparasite infecting wide ranges of marine organisms in coastal and oceanic waters. The parasitoid has received growing attention, due to its enormous genetic diversity in seawaters and suppressive effects on the growth of host dinoflagellates. Harmful algal blooms (HABs) caused by planktonic dinoflagellates have significantly impacted the coastal environment and mariculture in China. Series of studies have been conducted to reveal the occurrence mechanism and negative impacts of HABs in past decades, while the factors contributing to the recession of HABs have rarely been studied. Thus, the host range, prevalence and diversity of Amoebophrya along the coastline of China were systemically investigated to facilitate future studies on the ecological roles of the parasitoid. Overall, 10 dinoflagellate taxa were found to be infected by Amoebophrya spp., and the prevalence ranged from 0.03% to 2.50%. Sequencing of environmental genomic DNA revealed substantial diversity and significant regional heterogeneity of Amoebophryidae sequences derived from 12 coastal bays, while no significant correlation was observed among geographical locations. Phylogenetic analyses of 18S rDNA sequences derived from individual Amoebophrya-infected cells indicated the host divergence of the parasitoid and lend credence to the multiple species assumption. The results further revealed the broad host range, wide distribution and substantial diversity of Amoebophrya in the coastal waters of China, that should not be neglected in future studies on the succession of HABs, as well as the ecological significance of this parasitoid in marine microbial food webs.  相似文献   

7.
Phytoplankton blooms are a worldwide ecological problem and one of the major algae that cause phytoplankton blooms is Akashiwo sanguinea. Though much research has addressed the abiotic causes (e.g. growth condition) of A. sanguinea blooms, few studies have examined the dynamics of microbial communities associated with these blooms. In this study, polymerase chain reaction (PCR)-based denaturing gradient gel electrophoresis (DGGE) analysis of 16S rDNA genes was used to document changes in the phylogenetic diversity of microbial communities associated with an A. sanguinea bloom that occurred in the Xiamen sea in May 2010. Surface sea water was sampled once a day within five consecutive days at four sites, and the microbial community composition was determined using DGGE. Sea water concentrations of chlorophyll a, nitrate and phosphate were also measured. The results indicated that the A. sanguinea bloom was probably stimulated by low salinity (26–30‰) and ended probably because inorganic nutrients were consumed and resulted in a N/P ratio unfavorable for this alga. Gammaproteobacteria populations increased significantly during bloom declines and then decreased post-bloom. Divergences in the microbial community composition during different bloom periods were the result of changes in Candidatus, Pelagibacter, Alteromonas, Rhodobacteraceae, Vibrio and Pseudoalteromonas populations. Sediminimonas qiaohouensis was the first bacterium shown to be significantly negatively correlated with A. sanguinea concentration. This study indicated that bacteria may play an important role in A. sanguinea–bloom regulation and provides a deeper insight into bacterial community succession during and after an A. sanguinea–bloom.  相似文献   

8.
Mechanisms influencing initiation of harmful algal blooms (HABs) are diverse, and are not likely to be mutually exclusive. Rather, initiation of HABs is a result of interactions between processes, which result in biological, physical, and chemical conditions optimal for a bloom. Due to the complexity of some bloom initiation processes, bloom-preventative management may be possible. Results from a modeling exercise and a laboratory experiment indicated that a phytoplankton bloom could be circumvented through manipulation of the nutrient-loading mode, i.e., pulsed vs. continuous loading. These findings, should they prove consistent in more robust field experiments, may provide insights for the development of new management approaches for some HABs. Optimal bloom conditions, however, vary between HAB species. Consequently, it is unlikely that a single management solution will exist. Preventative management efforts will require early warning of HAB initiation, perhaps even before the appearance of an HAB species. An indicator based on the dynamic nature of phytoplankton succession events and phytoplankton species diversity may prove useful for this purpose. Applying this index to an existing plankton data set showed that Microcystis blooms might have been predicted months before the start of the bloom.  相似文献   

9.
Parasitic dinoflagellates of the genus Amoebophrya infect free-living dinoflagellates, some of which can cause harmful algal blooms (HABs). High prevalence of Amoebophrya spp. has been linked to the decline of some HABs in marine systems. The objective of this study was to evaluate the impact of Amoebophrya spp. on the dynamics of dinoflagellate blooms in Salt Pond (MA, USA), particularly the harmful species Alexandrium fundyense. The abundance of Amoebophrya life stages was estimated 3–7 days per week through the full duration of an annual A. fundyense bloom using fluorescence in situ hybridization coupled with tyramide signal amplification (FISH- TSA). More than 20 potential hosts were recorded including Dinophysis spp., Protoperidinium spp. and Gonyaulax spp., but the only dinoflagellate cells infected by Amoebophrya spp. during the sampling period were A. fundyense. Maximum A. fundyense concentration co-occurred with an increase of infected hosts, followed by a massive release of Amoebophrya dinospores in the water column. On average, Amoebophrya spp. infected and killed ∼30% of the A. fundyense population per day in the end phase of the bloom. The decline of the host A. fundyense population coincided with a dramatic life-cycle transition from vegetative division to sexual fusion. This transition occurred after maximum infected host concentrations and before peak infection percentages were observed, suggesting that most A. fundyense escaped parasite infection through sexual fusion. The results of this work highlight the importance of high frequency sampling of both parasite and host populations to accurately assess the impact of parasites on natural plankton assemblages.  相似文献   

10.
During the bloom events of the harmful dinoflagellate Cochlodinium polykrikoides in August and October, 2012, infections by two different Amoebophrya species were observed in Korean coastal waters. To investigate the dynamics of the two parasites and their relative impact on the host populations, a quantitative real-time PCR (qPCR) method was applied to detect and quantify the parasites in the free-living and parasitic stages. Each specific primer set of the target species, Amoebophrya sp. 1 and sp. 2 was designed on the large subunit (LSU) and the first internal transcribed spacer (ITS1) of ribosomal RNA (rRNA) gene, respectively. Dynamics of the two Amoebophrya species via qPCR assay showed distinct patterns during the C. polykrikoides bloom events. Amoebophrya sp. 1 showed peaks during both bloom events in August and October with relatively low copies (106 to 107 copies L−1), while Amoebophrya sp. 2 appeared only during the bloom event in October with very high copies (109 to 1010 copies L−1). Overall, the qPCR measurements for the dynamics of two Amoebophrya species in the parasitic stage (> 5 μm fractions) were consistent with parasite prevalence through microscopic observations. Amoebophrya sp. 1 infections were observed during both bloom events in August and October with relatively low parasite prevalence (0.1–1.5%), while Amoebophrya sp. 2 infections were detected only during the bloom event in October with high prevalence (up to 45%). Taken together, Amoebophrya sp. 1 may be a generalist and C. polykrikoides may not be its primary host, while Amoebophrya sp. 2 may be a specialist which can substantially impact host population dynamics.  相似文献   

11.
Sponges occur across diverse marine biomes and host internal microbial communities that can provide critical ecological functions. While strong patterns of host specificity have been observed consistently in sponge microbiomes, the precise ecological relationships between hosts and their symbiotic microbial communities remain to be fully delineated. In the current study, we investigate the relative roles of host population genetics and biogeography in structuring the microbial communities hosted by the excavating sponge Cliona delitrix. A total of 53 samples, previously used to demarcate the population genetic structure of C. delitrix, were selected from two locations in the Caribbean Sea and from eight locations across the reefs of Florida and the Bahamas. Microbial community diversity and composition were measured using Illumina‐based high‐throughput sequencing of the 16S rRNA V4 region and related to host population structure and geographic distribution. Most operational taxonomic units (OTUs) specific to Cliona delitrix microbiomes were rare, while other OTUs were shared with congeneric hosts. Across a large regional scale (>1,000 km), geographic distance was associated with considerable variability of the sponge microbiome, suggesting a distance–decay relationship, but little impact over smaller spatial scales (<300 km) was observed. Host population structure had a moderate effect on the structure of these microbial communities, regardless of geographic distance. These results support the interplay between geographic, environmental, and host factors as forces determining the community structure of microbiomes associated with C. delitrix. Moreover, these data suggest that the mechanisms of host regulation can be observed at the population genetic scale, prior to the onset of speciation.  相似文献   

12.
Cannibalism (CANN) and intraguild predation (IGP) may provide energy and nutrients to individuals and eliminate potential competitors. These negative competitive interactions could also affect the coexistence of predatory species. The co‐occurrence of aphidophagous ladybird species in crops creates opportunities for CANN and IGP, especially when aphids become scarce. The Lotka–Volterra model predicts the coexistence of two species if intraspecific competition is stronger than interspecific interference interactions. Cycloneda sanguinea L. and Eriopis connexa (Germar) (both Coleoptera: Coccinellidae) coexist in sweet pepper crops in La Plata (Argentina) consuming mainly Myzus persicae (Sulzer) (Hemiptera: Aphididae). The present study used laboratory experiments to estimate levels of CANN and IGP by adults and larvae on eggs, and by adults on larvae, in both the presence and absence of prey (i.e., M. persicae), to explain the effect of prey on coexistence of these two predators. Levels of CANN by C. sanguinea and E. connexa were high in the absence of aphids, and decreased when prey was present. Intraguild predation was bidirectional and asymmetric. Adults and larvae of E. connexa were more voracious IG predators of C. sanguinea than vice versa, the former being the stronger IG predator and interference competitor. Eriopis connexa always won when larvae of the same instar were compared, whereas the larger larva always won when larvae were of different instars, regardless of species. In the presence of prey, CANN by both species decreased, but IGP by E. connexa on C. sanguinea remained high, suggesting that E. connexa could displace C. sanguinea via interspecific interference competition. Other factors potentially affecting the coexistence of C. sanguinea and E. connexa in sweet pepper crops are discussed.  相似文献   

13.
Biotic interactions dominate plankton communities, yet the microbial consortia associated with harmful algal blooms (HABs) have not been well-described. Here, high-throughput amplicon sequencing of ribosomal genes was used to quantify the dynamics of bacterial (16S) and phytoplankton assemblages (18S) associated with blooms and cultures of two harmful algae, Alexandrium fundyense and Dinophysis acuminata. Experiments were performed to assess changes in natural bacterial and phytoplankton communities in response to the filtrate from cultures of these two harmful algae. Analysis of prokaryotic sequences from ecosystems, experiments, and cultures revealed statistically unique bacterial associations with each HAB. The dinoflagellate, Alexandrium, was strongly associated with multiple genera of Flavobacteria including Owenweeksia spp., Maribacter spp., and individuals within the NS5 marine group. While Flavobacteria also dominated Dinophysis-associated communities, the relative abundance of Alteromonadales bacteria strongly co-varied with Dinophysis abundances during blooms and Ulvibacter spp. (Flavobacteriales) and Arenicella spp. (Gammaproteobacteria) were associated with cells in culture. Eukaryotic sequencing facilitated the discovery of the endosymbiotic, parasitic dinoflagellate, Amoebophrya spp., that had not been regionally described but represented up to 17% of sequences during Alexandrium blooms. The presence of Alexandrium in field samples and in experiments significantly altered the relative abundances of bacterial and phytoplankton by both suppressing and promoting different taxa, while this effect was weaker in Dinophysis. Experiments specifically revealed a negative feedback loop during blooms whereby Alexandrium filtrate promoted the abundance of the parasite, Amoebophrya spp. Collectively, this study demonstrates that HABs formed by Alexandrium and Dinophysis harbor unique prokaryotic and eukaryotic microbiomes that are likely to, in turn, influence the dynamics of these HABs.  相似文献   

14.
This study examined a new method of mitigating harmful algal blooms (HABs) by combining biosurfactant sophorolipid and yellow clay. To investigate the effects and practicability of this HAB mitigation method, field experiments were carried out during a Cochlodinium bloom near Miruk Island, South Korea, in August 2002. Field experiments examined the effects of sophorolipid and yellow clay on Cochlodinium bloom mitigation and on marine plankton such as bacteriaplankton, heterotrophic protists, and zooplankton. A mixture of 5 mg l−1 sophorolipid and 1 g l−1 yellow clay was sprayed directly on the sea surface and its effect was compared with that of 10 g l−1 of yellow clay applied under similar conditions. The sophorolipid–yellow clay mixture more efficiently mitigated the Cochlodinium bloom (95% removal efficiency after 30 min) than yellow clay alone (79% after 30 min). Further, no variation in bacterial abundance occurred 30 min after spraying the sophorolipid–yellow clay mixture. After 30 min, heterotrophic protist abundance at the surface decreased 21 and 41%, respectively, following the sophorolipid–yellow clay mixture and yellow clay treatments. Zooplankton decreased by 38% 15 min after spraying the mixture and 67% 30 min after spraying the yellow clay. These results indicate that the mixture of sophorolipid and yellow clay had a less adverse effect on bacteriaplankton, heterotrophic protists, and zooplankton than the yellow clay, suggesting that the sophorolipid–yellow clay mixture can mitigate HABs efficiently with fewer negative effects on the pelagic ecosystem.  相似文献   

15.
Amoebophrya ceratii (Koeppen) Cachon is an obligate parasite of dinoflagellates and may represent a species complex. However, little is known about the biology and host range of different strains of Amoebophrya Cachon. Here, we determined parasite generation time and dinospore infectivity, survival, and ability to infect nonprimary hosts for strains of Amoebophrya from Akashiwo sanguinea (Hirasaka) G. Hansen et Moestrup, Gymnodinium instriatum (Freudenthal et Lee) Coats comb. nov., and Karlodinium micrum (Leadbeater et Dodge) J. Larsen. Akashiwo sanguinea was readily infected, with parasite prevalence reaching 100% in dinospore:host inoculations above a 10:1 ratio. Parasitism also approached 100% in G. instriatum, but only when inoculations exceeded a 40:1 ratio. Karlodinium micrum appeared partially resistant to infection, as parasite prevalence saturated at 92%. Parasite generation time differed markedly among Amoebophrya strains. Survival and infectivity of dinospores decreased over time, with strains from G. instriatum and A. sanguinea unable to initiate infections after 2 and 5 days, respectively. By contrast, dinospores from Amoebophrya parasitizing K. micrum remained infective for up to 11 days. Akashiwo sanguinea and G. instriatum were not infected when exposed to dinospores from nonprimary Amoebophrya strains. Karlodinium micrum, however, was attacked by dinospores of Amoebophrya from the other two host species, but infections failed to reach maturity. Observed differences in host–parasite biology support the hypothesis that Amoebophrya ceratii represents a complex of host‐specific species. Results also suggest that Amoebophrya strains have evolved somewhat divergent survival strategies that may encompass sexuality, heterotrophy during the “free‐living” dinospore stage, and dormancy.  相似文献   

16.
17.
18.
An algal bloom caused by the dinoflagellate Akashiwo sanguinea was observed in October–November 2009 along the central Oregon coast (44.6°N), off Newport, Oregon, U.S.A. In this paper, the conditions are described which led to the development and demise of this bloom. The bloom was observed for 1 month from 5-October until 4-November with the peak of abundance on 19-October (347,615 cells L−1). The A. sanguinea bloom followed September blooms of the diatoms Pseudo-nitzschia spp, Chaetoceros debilis, and the dinoflagellate Prorocentrum gracile. The bloom occurred when nitrate and silicate concentrations were <2 μM and <8 μM, respectively, and when the water column was stratified. This A. sanguinea dinoflagellate bloom event was closely related to the anomalous upwelling conditions in 2009: upwelling ceased early, at the end of August, whereas a normal upwelling continues into early October. This relaxation extended to near the end of September as a prolonged downwelling event, but then active upwelling reappeared in October and November. The explanation for the occurrence of the A. sanguinea bloom in October may be related to a combination of a prior diatom bloom, a stratified water column with low nutrient concentration in September, and an active upwelling event in October. As for the ultimate source of the cells, the hypothesis is that the seed stock for the A sanguinea bloom off Oregon was southward transport of cells from the Washington coast where a massive bloom of A. sanguinea was first observed in September 2009.  相似文献   

19.
Members of the Amoebophrya ceratii complex are endoparasitic dinoflagellates that parasitize a number of their dinoflagellate relatives, including toxic and/or harmful algal bloom-forming species. Despite many studies on the occurrence, prevalence, biology and molecular phylogeny of Amoebophrya spp., little attention has been given to toxin dynamics of host population following parasitism. Using Amoebophrya sp. infecting the paralytic shellfish toxin (PSP)-producing dinoflagellate Alexandrium fundyense, we addressed the following questions: (1) does parasitism by Amoebophrya sp. alter toxin content and toxin profiles of the dinoflagellate A. fundyense over the infection cycle? and (2) do parasite dinospores produced at the end of the infection cycle retain host toxins and thus potentially act as a vector to convey PSP toxin through the marine microbial food-web? Toxin time-course experiments showed that the PSP toxin contents did not vary significantly over the infection cycle, but mean toxin content for infected cultures was significantly higher than that for uninfected cultures. Host toxins were not detected in the free-living, dinospore stage of the parasite. Therefore, our results indicate that Amoebophrya sp. does not function as a vector for transferring PSP toxins to higher trophic levels. Rather, Amoebophrya infections appear to play an important role in maintaining healthy ecosystems by transforming potent toxins-producing dinoflagellates into non-toxic dinospores, representing “edible food” for consumers of the marine microbial food-web during toxic algal bloom event.  相似文献   

20.
Harmful algal blooms (HABs) are globally expanding, compromising water quality worldwide. HAB dynamics are determined by a complex interplay of abiotic and biotic factors, and their emergence has often been linked to eutrophication, and more recently to climate change. The dinoflagellate Alexandrium is one of the most widespread HAB genera and its success is based on key functional traits like allelopathy, mixotrophy, cyst formation and nutrient retrieval migrations. Since 2012, dense Alexandrium ostenfeldii blooms (up to 4500 cells mL−1) have recurred annually in a creek located in the southwest of the Netherlands, an area characterized by intense agriculture and aquaculture. We investigated how physical, chemical and biological factors influenced A. ostenfeldii bloom dynamics over three consecutive years (2013–2015). Overall, we found a decrease in the magnitude of the bloom over the years that could largely be linked to changing weather conditions during summer. More specifically, low salinities due to excessive rainfall and increased wind speed corresponded to a delayed A. ostenfeldii bloom with reduced population densities in 2015. Within each year, highest population densities generally corresponded to high temperatures, low DIN:DIP ratios and low grazer densities. Together, our results demonstrate an important role of nutrient availability, absence of grazing, and particularly of the physical environment on the magnitude and duration of A. ostenfeldii blooms. Our results suggest that predicted changes in the physical environment may enhance bloom development in future coastal waters and embayments.  相似文献   

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