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Marine phytoplankton communities are pivotal in biogeochemical cycles and impact global climate change. However, the dynamics of the dinoflagellate community, its co-occurrence relationship with other eukaryotic plankton communities, and environmental factors remain poorly understood. In this study, we aimed to analyze the temporal changes in the eukaryotic plankton community using a 18S rDNA metabarcoding approach. We performed intensive monitoring for 439 days at intervals of three days during the period from November 2018 to June 2020 (n = 260) in Jangmok Bay Time-series Monitoring Site in South Korea. Among the 16,224 amplicon sequence variants (ASVs) obtained, dinoflagellates were the most abundant in the plankton community (38 % of total relative abundance). The dinoflagellate community was divided into 21 groups via cluster analysis, which showed an annually similar distribution of low-temperature periods. Additionally, we selected 11 taxa that had an occurrence mean exceeding 1 % of the total dinoflagellate abundance, accounting for 93 % of the total dinoflagellate community: namely Heterocapsa rotundata, Gymnodinium sp., Akashiwo sanguinea, Amoebophrya sp., Euduboscquella sp., Spiniferites ramosus, Dissodinium pseudolunula, Sinophysis sp., Karlodinium veneficum, and Katodinium glaucum. The key dinoflagellate species were well represented at temporally variable levels over an entire year. Heterocapsa rotundata was not significantly affected by water temperature, whereas its dynamics were largely influenced by strong predation pressure, competition, and/or the supplementation of food sources. The growth of A. sanguinea was associated with dissolved inorganic phosphorus concentrations, while Euduboscquella sp. showed a significant relationship with D. pseudolunula and K. glaucum, largely representing a positive association that implies possible parasitic mechanisms. This study demonstrated interactions between key dinoflagellate species and the environment, as well as parasites, predators, competitors, and feeders.

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The production of allelochemicals by the toxigenic dinoflagellate Alexandrium catenella is one of the suggested mechanisms to facilitate its bloom formation and persistence by outcompeting other phototrophic protists and reducing grazing pressure. In Southern California, toxic events caused by A. catenella and paralytic shellfish toxins (PSTs) regularly impact coastal ecosystems; however, the trophic interactions and mechanisms promoting this species in a food web context are still not fully understood. In the present study, we combined a dynamical mathematical model with laboratory experiments to investigate potential toxic and allelochemical effects of an A. catenella strain isolated off the coast of Los Angeles, Southern California, on competitors and a common zooplankton consumer. Experiments were conducted using three toxigenic strains of A. catenella, comparing the new Californian isolate (Alex Cal) to two strains previously described from the North Sea, a lytic (Alex2) and non-lytic (Alex5) strain, testing for donor density-dependent effects on two phytoplankton species (Rhodomonas salina, Tetraselmis sp.) and on the rotifer Brachionus plicatilis. Bioassays revealed a steep decline in competitor and consumer populations with increasing Alex Cal concentrations, indicating an intermediate lytic activity compared to the North Sea strains (lytic Alex2 and non-lytic Alex5). The rotifer fed and grew well on the PST- toxic, but non-lytic Alex5 strain, while its survival significantly decreased with increasing concentrations of the two lytic strains Alex Cal and Alex 2, indicating that negative effects on the rotifer were mediated by allelochemicals rather than PST-toxins. Mixed culture experiments including both competitors and consumers demonstrated that the intensity of allelochemical effects not only depended on the A. catenella density but also on the target density. Negative effects on grazers were alleviated by co-occurring competitors with a lower sensitivity to allelochemicals, thus reducing harmful compounds and allowing grazing control on the dinoflagellate to come into effect again. Results from mixed culture experiments were supported by the mathematical approach used in this study which was calibrated with data from simple monoculture growth, pairwise competition and predator-prey experiments, demonstrating the applicability of this model approach to predict the outcome of more complex food web dynamics at the community level.

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Studies on dinoflagellate cysts in the Arabian Sea (AS) are limited to the coastal waters, but no information from the deeper depths. The dinoflagellate cyst assemblages in surface sediment samples (0-2 cm) from the deeper depths (up to ~ 4500 m) of central (oxygen minimum zone (OMZ)) and southeastern (oligotrophic) AS revealed that the relatively good numbers of cyst concentrations reach deeper depths of OMZ (3505 m) and oligotrophic (4368 m) regions, but the former harbored more cyst concentrations than the latter. The cyst concentration and species count (including HAB species) recorded here are lower compared to the eastern (EAS) and western (WAS) AS, but the autotrophic cyst dominance (74-83%) at deeper depths is in contrast with the heterotrophic dominance in coastal AS. Of the recorded 41 cyst species (belonging to 18 genera), four species (cyst of Cladopyxis sp., Operculodinium janduchenei, Stelladinium bifurcatum, and Protoperidinium monospinum) from the deepest part of oligotrophic AS form the first report. In contrast, Spiniferites and Lingulodinium cysts were common occurrences. Taxonomic comparison with literature revealed (i) the prevalence of more cosmopolitan species (32 species) which could be due to the prevalence of large and small-scale lateral transport of cysts in oligotrophic regions followed by OMZ and coastal regions, respectively, and (ii) very few region-specific species, i.e., cyst of Protoperidinium latissimum, Lejeunecysta sabrina, cyst of Protoperidinium denticulatum in EAS and Impagidinium patulum, and I. strialatum, in WAS. Interestingly, variability in the morphometry was evident between the coastal and open oceans in some cosmopolitan cysts, e.g., Operculodinium centrocarpum and Lingulodinium machaerophorum. These findings from the less studied pelagic regions will contribute to the growing knowledge of dinoflagellate cyst distribution patterns and highlight the significance of cyst taxa and morphology as potential ecological indicators for AS.

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Ship ballast residual sediments are an important vector of introduction for non-indigenous species. We evaluated the proportion of residual sediments and associated organisms released during de-ballasting operations of a commercial bulk carrier and estimated a total residual sediment accumulation of ∼13 t, with accumulations of up to 20 cm in some tank areas that had accumulated over 11 years. We observed interior hull-fouling (anemones, hydrozoans, and bryozoans) and high abundances of viable invertebrate resting stages and dinoflagellate cysts in sediments. Although we determined that <1 % of residual sediments and associated resting stages were resuspended and released into the environment during individual de-ballasting events, this represents a substantial inoculum of 21 × 107 viable dinoflagellate cysts and 7.5 × 105 invertebrate resting stages with many taxa being nonindigenous, cryptogenic, or toxic/harmful species. The methods used and results will help estimate propagule pressure associated with this pathway and will be relevant for residual sediments and nonindigenous species management.

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Variation in the condition of marine sediments provides selective preservation milieus, which act as a key determinant for the abundance and distribution of dinoflagellate resting cysts in natural sediments. Microbial degradation is an understudied biological factor of potential importance in the processes. However, gaps remain in our knowledge about the fundamental information of the bacterial consortia associated with dinoflagellate resting cysts both in laboratory cultures and in the field. Here we used Scrippsiella acuminata as a representative of cyst-producing dinoflagellates to delineate the diversity and composition of bacterial microbiomes co-existing with the laboratory-cultured resting cysts, and to explore possible impacts of low temperature, darkness, and anoxia (the mock conditions commonly observed in marine sediments) on the associated bacterial consortia. Bacterial microbiome with high diversity were revealed associated with S. acuminata at resting stage. The mock conditions could significantly shift bacterial community structure and exert notably inhibitory effects on growth-promoting bacteria. Resting cysts under conditions typically observed in marine sediments fostered bacterial microbiomes with more diverse trophic strategies, characteristic of prominently enriched anaerobic chemotrophic bacteria generating energy via respiration with several different terminal electron acceptors, which yielded more acidic milieu unfavorable for the preservation of calcareous resting cysts. Our findings suggest that there is complex and dynamic interaction between dinoflagellates resting cysts and the associated bacterial consortia in natural sediments. This intrinsic interaction may influence the maintenance and/or accumulation of dinoflagellate resting cysts with potential of germination and initiation blooms in the field.

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Dinoflagellates are one of the largest groups of marine microalgae and exhibit diverse trophic strategies. Some dinoflagellates can produce secondary metabolites that are known to be toxic, which can lead to ecologically harmful blooms. Amphidinium carterae is one species of dinoflagellate that produces toxic compounds and is used as a model for dinoflagellate studies. The impact of the microbiome on A. carterae growth and metabolite synthesis is not yet fully understood, nor is the impact of bacterial data on sequencing and assembly. An antibiotic cocktail was previously shown to eliminate 16S amplification from the dinoflagellate culture. Even with drastically reduced bacterial numbers during antibiotic treatment, bacterial sequences were still present. In this experiment, we used novel Nanopore long-read sequencing techniques on A. carterae cultures to assemble 15 full bacterial genomes ranging from 2.9 to 6.0 Mb and found that the use of antibiotics decreased the percentage of reads mapping back to bacteria. We also identified shifts in the microbiome composition and identified a potentially deleterious bacterial species arising in the absence of the antibiotic treatment. Multiple antibiotic resistance genes were identified, as well as evidence that the bacterial population does not contribute to toxic secondary metabolite synthesis.

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Paralytic shellfish toxins (PSTs) are potent neurotoxins produced by certain microalgae, particularly dinoflagellates, and they can accumulate in shellfish in coastal seawater and thus pose significant health risks to humans. To explore the relationship between toxicity and PST profiles in seawater and mussels, the spatiotemporal variations in PST concentrations and profiles were investigated along the southern coast of Korea under peak PST levels during spring. Seawater and mussel samples were collected biweekly from multiple stations, and the toxin concentrations in the samples were measured. Moreover, the dinoflagellate community composition was analyzed using next-generation sequencing to identify potential PST-producing species. The PST concentrations and toxin profiles showed substantial spatiotemporal variability, with GTX1 and GTX4 representing the dominant toxins in both samples, and C1/2 tending to be higher in seawater. Alexandrium species were identified as the primary sources of PSTs. Environmental factors such as water temperature and salinity influenced PST production. This study demonstrates that variability in the amount and composition of PSTs is due to intricate ecological interactions. To mitigate shellfish poisoning, continuous monitoring must be conducted to gain a deeper understanding of these interactions.

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In the coastal waters around Shandong peninsula, an unprecedented winter bloom of dinoflagellates Gonyaulax polygramma and Akashiwo sanguinea occurred in 2021 from late November to early December. The bloom affected a wide area of coastal waters extending from west to east along the northern Shandong peninsula and had a devastating blow to the kelp cultivation industry. Based on the remote-sensing data, the initiation of the bloom was traced back to the region adjacent to the mouth of the Yellow River in Laizhou Bay, where enhanced freshwater discharge from the Yellow River was recorded from September to November. It's proposed that the increased precipitation in the Yellow River basin associated with northward extension of the precipitation band in China could be an important reason for this winter bloom. This unusual winter bloom around Shandong peninsula highlights the potential risks of harmful algal blooms and their impacts on coastal ecosystems under the background of climate change.

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Dakhla Bay, situated on the African Atlantic coast, has witnessed sporadic harmful algal blooms (HABs) caused by toxic dinoflagellate species over the past two decades. In this study, we investigated the distribution, abundance, and diversity of dinoflagellate cysts, with a focus on potentially toxic species that develop in this ecosystem where such data are lacking. Sediment samples were collected in April 2018 through coring at 49 stations distributed across the bay. The highest abundance of dinoflagellate cysts was recorded at 304 cysts/g dry sediment, observed at the inner part of the bay, indicating that this area is the preferential zone for cyst accumulation. Pearson's tests revealed significant positive correlations (P < 0.05) between cyst abundance and the water content, organic matter, and fine fraction (< 63 µm) of the sediment. Cyst morphotypes of potentially toxic dinoflagellate species known to produce saxitoxins, such as Alexandrium minutum, Alexandrium tamarense species complex, Gymnodinium catenatum, and yessotoxins, such as Lingulodinium polyedrum and Gonyaulax cf. spinifera, were identified in the sediment of Dakhla Bay. These findings were further supported by our long-term monitoring period (2005-2018), underscoring the presence of these HAB species in Dakhla Bay. During our survey, sporadic mollusk intoxication events were recorded at station PK25 for the grooved razor shell Solen marginatus and at station Boutelha for the oyster Crassostrea gigas. Paralytic shellfish toxin concentrations exceeded the sanitary threshold (80 µg STX di-HCl eq/100 g of shellfish meat) only twice, in December 2006 and January 2007 at station PK25. Contamination by amnesic shellfish toxins occurred sporadically but never reached the sanitary threshold of 20 µg/g of shellfish meat. Lipophilic shellfish intoxication occurred multiple times in the two investigated areas. These observations suggest that the cysts of the identified HAB species germinated and inoculated the water column, resulting in the observed intoxication events. Relatively low levels of intoxication could be attributed to the moderate abundances of cysts, which may reduce the seeding capacity. This could be explained by the significant interaction of Dakhla Bay with the Atlantic Ocean, characterized by hydrological dynamics that impede the deposition and accumulation of cysts in the bay's sediments. This study reaffirms the importance of investigating dinoflagellate cysts in assessing the diversity of HAB species and evaluating associated sanitary risks.

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The aquaculture industry requires green solutions to solve several environmental challenges, including adequate wastewater remediation and natural drug applications to treat bacteria- and virus-related diseases. This study investigated the feasibility of cultivating the dinoflagellate Durusdinium glynnii in aquaculture wastewater from shrimp rearing in a synbiotic system (AWW-SS), with different dilutions of f/2 medium (FM). Interestingly, D. glynnii demonstrated enhanced growth in all AWW-SS treatments compared to the control (FM). The highest growth rates were achieved at AWW-SS:FM dilutions of 75:25 and 50:50. The removal of total nitrogen and total phosphorus reached 50.1 and 71.7%, respectively, of the crude AWW-SS. Biomass extracts of D. glynnii grown with AWW-SS were able to inhibit the growth of the bacteria Vibrio parahaemolyticus (inhibition zone of 10.0 ± 1.7 mm) and V. vulnificus (inhibition zone of 11.7 ± 1.5 mm). The presented results demonstrate that the dinoflagellate D. glynnii is a potential candidate for the development of circularity for sustainable aquaculture production, particularly by producing anti-Vibrio compounds at a near-zero cost.

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Dinophysis, a mixotrophic dinoflagellate that is known to prey on the ciliate Mesodinium rubrum, and retain its chloroplasts, is responsible for diarrhetic shellfish poisoning (DSP) in humans and has been identified on all U.S. coasts. Monocultures of Dinophysis have been used to investigate the growth of Dinophysis species in response to variations in environmental conditions, however, little is known about the roles of system stability (turbulence) and mixotrophy in the growth and toxicity of Dinophysis species in the U.S.. To begin to address this gap in knowledge, culturing experiments were conducted with three species (four strains) of Dinophysis, that included predator-prey co-incubation (Dinophysis spp.+ M. rubrum) and prey-only (M. rubrum) flasks. Cultures were investigated for effects of low or high turbulence on Dinophysis spp. growth, feeding, and amounts of intra- and extracellular toxins: okadaic acid and derivatives (diarrhetic shellfish toxins, DSTs) and pectenotoxins (PTXs). Turbulence did not have a measurable effect on the rates of ingestion of M. rubrum prey by Dinophysis spp. for any of the four strains, however, effects on growth and particulate and dissolved toxins were observed. High turbulence (ε = 10-2 m2s-3) significantly slowed growth of both D. acuminata and D. ovum relative to still controls, but significantly stimulated growth of the D. caudata strain. Increasing turbulence also resulted in significantly higher intracellular toxin content in D. acuminata cultures (DSTs and PTXs), but significantly reduced intracellular toxin content (PTXs) in those of D. caudata. An increase in turbulence appeared to promote toxin leakage, as D. ovum had significantly more extracellular DSTs found in the medium under high turbulence when compared to the still control. Overall, significant responses to turbulence were observed, whereby the three strains from the "Dinophysis acuminata complex" displayed a stress response to turbulence, i.e., decreasing growth, increasing intracellular toxin content and/or increasing toxin leakage, while the D. caudata strain had an opposite response, appearing stimulated by, or more tolerant of, high turbulence.

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We analyzed surface sediments from 12 stations located in Izmir Bay to determine the impact of anthropogenic pollution on dinoflagellate cysts. Forty-three dinoflagellate cyst taxa and two cyst assemblage zones were identified. Zone 1 is characterized by the dominance of cysts of Gymnodinium nolleri, Selenopemphix nephroides, and Operculodinium centrocarpum from the oligotrophic part of Izmir Bay. Zone 2 is in the highly productive inner part of the bay and is characterized by the high abundances of Lingulodinium machaerophorum, Spiniferites ramosus, cysts of Scrippsiella spp., cysts of Polykrikos spp. and Quinquecuspis concreta. We used multivariate statistical analysis (DCA and CCA) on dinoflagellate cysts and environmental variables to support the identification of Zones 1 and 2. Our analyses also revealed that summer and winter chlorophyll-a as well as elevated nitrate and nitrite concentrations are significant parameters in controlling dinoflagellate cyst distribution in Izmir Bay.

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Dinoflagellates are an abundant and diverse group of protists representing a wealth of unique biology and ecology. While many dinoflagellates are photosynthetic or mixotrophic, many taxa are heterotrophs, often with complex feeding strategies. Compared to their photosynthetic counterparts, heterotrophic dinoflagellates remain understudied, as they are difficult to culture. One exception, a long-cultured isolate originally classified as Amphidinium but recently reclassified as Oxytoxum, has been the subject of a number of feeding, growth, and chemosensory studies. This lineage was recently determined to be closely related to Prorocentrum using phylogenetics of ribosomal RNA gene sequences, but the exact nature of this relationship remains unresolved. Using transcriptomes sequenced from culture and three single cells from the environment, we produce a robust phylogeny of 242 genes, revealing Oxytoxum is likely sister to the Prorocentrum clade, rather than nested within it. Molecular investigations uncover evidence of a reduced, nonphotosynthetic plastid and proteorhodopsin, a photoactive proton pump acquired horizontally from bacteria. We describe the ultrastructure of O. lohmannii, including densely packed trichocysts, and a new type of mucocyst. We observe that O. lohmannii feeds preferentially on cryptophytes using myzocytosis, but can also feed on various phytoflagellates using conventional phagocytosis. O. lohmannii is amenable to culture, providing an opportunity to better study heterotrophic dinoflagellate biology and feeding ecology.

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The shift between photoautotrophic and phagotrophic strategies in mixoplankton significantly impacts the planktonic food webs and biogeochemical cycling. Considering the projected global warming, studying how temperature impacts this shift is crucial. Here, we combined the transcriptome of in-lab cultures (mixotrophic dinoflagellate Lepidodinium sp.) and the metatranscriptome dataset of the global ocean to investigate the mechanisms underlying the shift of trophic strategies and its relationship with increasing temperatures. Our results showed that phagocytosis-related pathways, including focal adhesion, regulation of actin cytoskeleton, and oxidative phosphorylation, were significantly stimulated in Lepidodinium sp. when cryptophyte prey were added. We further compared the expression profiles of photosynthesis and phagocytosis genes in Lepidodinium sp. in the global sunlit ocean. Our results indicated that Lepidodinium sp. became more phagotrophic with increasing temperatures when the ambient chlorophyll concentration was >0.3 mg.m-3 (~20.58% of the ocean surface) but became more photoautotrophic with increasing temperatures when the chlorophyll concentration was between 0.2 and 0.3 mg.m-3 (~11.47% of the ocean surface). Overall, we emphasized the crucial role of phagocytosis in phago-mixotrophy and suggested that the expression profile of phagocytosis genes can be a molecular marker to target the phagotrophic activity of mixoplankton in situ.

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Ships' ballast water and sediments have long been linked to the global transport and expansion of invasive species and thus have become a hot research topic and administrative challenge in the past decades. The relevant concerns, however, have been mainly about the ocean-to-ocean invasion and sampling practices have been almost exclusively conducted onboard. We examined and compared the dinoflagellate cysts assemblages in 49 sediment samples collected from ballast tanks of international and domestic routes ships, washing basins associated with a ship-repair yard, Jiangyin Port (PS), and the nearby area of Yangtze River (YR) during 2017-2018. A total of 43 dinoflagellates were fully identified to species level by metabarcoding, single-cyst PCR-based sequencing, cyst germination and phylogenetic analyses, including 12 species never reported from waters of China, 14 HABs-causing, 9 toxic, and 10 not strictly marine species. Our metabarcoding and single-cyst sequencing also detected many OTUs and cysts of dinoflagellates that could not be fully identified, indicating ballast tank sediments being a risky repository of currently unrecognizable invasive species. Particularly important, 10 brackish and fresh water species of dinoflagellate cysts (such as Tyrannodinium edax) were detected from the transoceanic ships, indicating these species may function as alien species potentially invading the inland rivers and adjacent lakes if these ships conduct deballast and other practices in fresh waterbodies. Significantly higher numbers of reads and OTUs of dinoflagellates in the ballast tanks and washing basins than that in PS and YR indicate a risk of releasing cysts by ships and the associated ship-repair yards to the surrounding waters. Phylogenetic analyses revealed high intra-species genetic diversity for multiple cyst species from different ballast tanks. Our work provides novel insights into the risk of bio-invasion to fresh waters conveyed in ship's ballast tank sediments and washing basins of shipyards.

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Reports of the benthic dinoflagellate Ostreopsis spp. have been increasing in the last decades, especially in temperate areas. In a context of global warming, evidences of the effects of increasing sea temperatures on its physiology and its distribution are still lacking and need to be investigated. In this study, the influence of temperature on growth, ecophysiology and toxicity was assessed for several strains of O. cf. siamensis from the Bay of Biscay (NE Atlantic) and O. cf. ovata from NW Mediterranean Sea. Cultures were acclimated to temperatures ranging from 14.5 °C to 32 °C in order to study the whole range of each strain-specific thermal niche. Acclimation was successful for temperatures ranging from 14.5 °C to 25 °C for O. cf. siamensis and from 19 °C to 32 °C for O. cf. ovata, with the highest growth rates measured at 22 °C (0.54-1.06 d-1) and 28 °C (0.52-0.75 d-1), respectively. The analysis of cellular content of pigments and lipids revealed some aspects of thermal acclimation processes in Ostreopsis cells. Specific capacities of O. cf. siamensis to cope with stress of cold temperatures were linked with the activation of a xanthophyll cycle based on diadinoxanthin. Lipids (neutral reserve lipids and polar ones) also revealed species-specific variations, with increases in cellular content noted under extreme temperature conditions. Variations in toxicity were assessed through the Artemia franciscana bioassay. For both species, a decrease in toxicity was observed when temperature dropped under the optimal temperature for growth. No PLTX-like compounds were detected in O. cf. siamensis strains. Thus, the main part of the lethal effect observed on A. franciscana was dependent on currently unknown compounds. From a multiclonal approach, this work allowed for defining specificities in the thermal niche and acclimation strategies of O. cf. siamensis and O. cf. ovata towards temperature. Potential impacts of climate change on the toxic risk associated with Ostreopsis blooms in both NW Mediterranean Sea and NE Atlantic coast is further discussed, taking into account variations in the geographic distribution, growth abilities and toxicity of each species.

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The dinoflagellate cysts present in the ballast water sediment of foreign ships in Shanghai Port have not been previously studied. Therefore, sediment samples were collected from the ballast water of 16 foreign ships in Shanghai Port, and the types of dinoflagellate cysts were identified and their abundance was calculated, with a specific focus on the analysis of toxic and harmful dinoflagellates. Moreover, simulations of temperature and salinity conditions throughout the year in the Shanghai port waters were conducted to carry out dinoflagellate cyst germination experiments, with analyze and compare the germinated dinoflagellate cysts under different conditions. Dinoflagellate cysts were found in 100 % of the ship sediment samples, including a total of 9 species of toxic and harmful dinoflagellate cysts. In the germination experiment, 15 °C was found to be the optimal temperature for the germination of dinoflagellate cysts in ballast water sediment, and high salinity is more favorable for cyst germination.

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Carbohydrates play a pivotal role in nutrient recycling and regulation of algal-bacterial interactions. Despite their ecological significance, the intricate molecular mechanisms governing regulation of phycosphere carbohydrates by bacterial taxa linked with natural algal bloom have yet to be fully elucidated. Here, a comprehensive temporal metagenomic analysis was conducted to explore the carbohydrate-active enzyme (CAZyme) genes in two discrete algal bloom microorganisms (Gymnodinium catenatum and Phaeocystis globosa) across three distinct bloom stages: pre-bloom, peak bloom, and post-bloom. Elevated levels of extracellular carbohydrates, primarily rhamnose, galactose, glucose, and arabinose, were observed during the initial and post-peak stages. The prominent CAZyme families identified-glycoside hydrolases (GH) and carbohydrate-binding modules (CBMs)-were present in both algal bloom occurrences. In the G. catenatum bloom, GH23/24 and CBM13/14 were prevalent during the pre-bloom and peak bloom stages, whereas GH2/3/30 and CBM12/24 exhibited increased prevalence during the post-bloom phase. In contrast, the P. globosa bloom had a dominance of GH13/23 and CBM19 in the initial phase, and this was succeeded by GH3/19/24/30 and CBM54 in the later stages. This gene pool variation-observed distinctly in specific genera-highlighted the dynamic structural shifts in functional resources driven by temporal alterations in available substrates. Additionally, ecological linkage analysis underscored a correlation between carbohydrates (or their related genes) and phycospheric bacteria, hinting at a pattern of bottom-up control. These findings contribute to understanding of the dynamic nature of CAZymes, emphasizing the substantial influence of substrate availability on the metabolic capabilities of algal symbiotic bacteria, especially in terms of carbohydrates.

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Our understanding of dinoflagellates' present-day and past ecology is limited due to the scarcity of data on the transport of dinoflagellate cysts in oceanic environments. Previous studies have shown that lateral transport affects the source-to-sink trajectory of cysts in the very productive region off Cape Blanc (NW Africa). Unsolved questions remain, such as: how far these cysts can be advected, whether the cyst sources vary over time and whether lateral transport is a permanent feature or restricted to individual events. To fill these gaps and assess the role of nepheloid layers on the lateral transport and preservation of dinoflagellate cysts, new data on dinoflagellate cyst distributions in the water column and sediments along a land-sea transect were obtained. Samples were collected in November 2018 along a shelf break-offshore transect during intense upwelling, notably, within and between the nepheloid layers. The composition and abundance of cysts with organic walls in the water column and surface sediments were studied. Moreover, the distribution of calcareous cysts was also analysed in the water samples, using non-destructive acid-free preparation methods. The records were dominated by empty cysts, but no clear indications that these originated from local resuspension of older sediments were observed. Clustering, principal component analysis and redundant discriminant analysis were used to compare cyst assemblages in the water column and surface sediments, and environmental conditions in the upper water column. The strong similarity in species composition of water samples collected in the active upwelling region to those collected from the more onshore parts of the Benthic Nepheloid Layer (BNL), upper Intermediate Nepheloid Layer (INL) (∼1000 m depth) and lower INL (∼2200 m depth) indicated that lateral transport of cysts within these NLs occurred until about ∼110 km from the shelf break. Cyst assemblages from above and below these NLs showed significantly different taxa composition reinforcing the role of NLs in the lateral advection of cysts. In the more offshore stations, vertically similar cyst assemblages were observed in the same station, independent of the sample depth, within or between the NLs, which supported that at these stations vertical transport was the dominant process influencing cyst assemblages. Consequently, the cyst signal in sediments off Cape Blanc may be affected both by horizontal transport of allochthonous cysts and vertical deposition of locally-produced cysts, particularly in the more offshore stations (>2000 m depth). Despite lateral transport and possible species-specific preservation effects, horizontal distributions of most cyst taxa in the water column and the surface sediments could be explained to a great extent by the main environmental gradients in the upper water column. This agrees with observations made in other regions, and reinforces that dinoflagellate cysts as good proxies to reconstruct past environmental conditions in offshore environments. New data on dinoflagellate cyst distribution, transport and accumulation patterns in deep environments off Cape Blanc may be useful for interpreting past environmental signals in the region. This is particularly relevant regarding calcareous cysts, as information on their distribution and ecology is very scarce. The present work contributes to a better understanding of the dispersal patterns of dinoflagellate cysts in the deep ocean, highlighting the significant role played by nepheloid layers in this process and thus on the dinoflagellate cyst signature in deep-sea sediments.

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Bioluminescence is light chemically produced by an organism. It is widespread across all major marine phyla and has evolved multiple times, resulting in a high diversity of spectral properties and first flash kinetic parameters (FFKP). The bioluminescence of a system is often a good proxy for planktonic biomass. The species-specific parameters of bioluminescent displays can be measured to identify species in situ and describe planktonic biodiversity. Most bioluminescent organisms will flash when mechanically stimulated i.e., when subjected to supra-threshold levels of shear stress. Here we compare first flash kinetic parameters such as flash duration, peak intensity, rise time, decay time, first-flash mechanically stimulated light and e-folding time obtained with the commercially available Underwater Bioluminescence Assessment Tool (UBAT). We provide descriptions of the first flash kinetic parameters of several species of dinoflagellates Pyrocystis fusiformis, Pyrocystis noctiluca, Pyrodinium bahamense, Lingulodinium polyedra, Alexandrium monilatum and two zooplankton (the ctenophore Mnemiopsis leidyi and the larvacean Oikopleura sp.). FFKPs are then compared and discussed using non-parametric analyses of variance (ANOVAs), hierarchical clustering and a linear discriminant analysis to assess the ability to use bioluminescence signatures for identification. Once the first flash kinetic parameters of a bioluminescent species have been described, it is possible to detect its presence using emissions collected by in situ bathyphotometers. Assessing abundance and diversity of bioluminescent species may therefore be possible.

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