Spatial, seasonal and species variations of harmful algal blooms in the South Yellow Sea and East China Sea

The occurrences of harmful algal blooms (HABs), in terms of frequency and area in the Chinese coastal waters, have been increasing since 1980s and caused considerable economic losses. In the present study, we have analyzed spatial and seasonal characteristics of HAB events in the southern Yellow Sea and East China Sea along Chinese coast from 1933 to 2004. With a total 435 HAB records, the most frequent HAB occurrence area (FHA) is off the Yangtze River mouth and another two FHA areas are located south of the Yangtze River estuary along about isobaths of 30–60 m coastal water in the East China Sea. The time of HAB occurrence shifted during our study period: from autumn (August–October) before 1980s to July–August in 1980s, during May–July in 1990s, and May–June for the period of 2000–2004. Causative species were found to be different: Noctiluca scintillans and Skeletonema costatum were dominant causative species prior to 2000; and Prorocentrum donghaiense Lu was dominant from 2000 to 2004 and also caused large blooms in May. Trichodesmium sp. caused many HABs in autumn (August–October) prior to 1980s with only one HAB between 1980 and 2004. The changes of the dominant HAB species may have affected the timings of HAB occurrence, as well as the increasing HAB-affected areas in recent years.     

Occurrences of harmful algal blooms (HABs) associated with ocean environments in the South China Sea

Harmful algal blooms (HABs) occur frequently in the South China Sea (SCS), causing enormous economic losses in aquaculture. We analyzed historical HAB records during the period from 1980 to 2003 in SCS. We found that HABs-affected areas have expanded and the frequency of HABs varied during this period. The seasonal and annual variations, as well as causative algal species of HABs are different among the four regions. Areas with frequent HABs include the Pearl River Estuary (China), the Manila Bay (the Philippines), the Masinloc Bay (the Philippines), and the western coast of Sabah (Malaysia). HABs occurred frequently during March–May in the northern region of SCS, May–July in the eastern region, July in the western region, and year-round in the southern region. Among the species that cause HABs, Noctiluca scintillans dominated in the northern region, and Pyrodinium bahamense in the southern and eastern regions. Causative species also varied in different years for the entire SCS. Both P. bahamense and N. scintillans were the dominant species during 1980–2003. Some species not previously recorded formed blooms during 1991–2003, including Phaeocystis globosa, Scrippsiella trochoidea, Heterosigma akashiwo, and Mesodinium rubrum. Variations in HABs are related to various regional conditions, such as a reversed monsoon wind in the entire SCS, river discharges in the northern area, upwelling in Vietnam coastal waters during southwest winds and near Malaysia coastal waters during northeast winds, and eutrophication from coastal aquaculture in the Pearl River estuary, Manila Bay, and Masinloc Bay. Handling editor: D. Hamilton     

In vitro interactions between several species of harmful algae and haemocytes of bivalve molluscs

Harmful algal blooms (HABs) can have both lethal and sublethal impacts on shellfish. To understand the possible roles of haemocytes in bivalve immune responses to HABs and how the algae are affected by these cells (haemocytes), in vitro tests between cultured harmful algal species and haemocytes of the northern quahog (= hard clam) Mercenaria mercenaria, the soft-shell clam Mya arenaria, the eastern and Pacific oysters Crassostrea virginica and Crassostrea gigas and the Manila clam Ruditapes philippinarum were carried out. Within their respective ranges of distribution, these shellfish species can experience blooms of several HAB species, including Prorocentrum minimum, Heterosigma akashiwo, Alexandrium fundyense, Alexandrium minutum and Karenia spp.; thus, these algal species were chosen for testing. Possible differences in haemocyte variables attributable to harmful algae and also effects of haemolymph and haemocytes on the algae themselves were measured. Using microscopic and flow cytometric observations, changes were measured in haemocytes, including cell morphology, mortality, phagocytosis, adhesion and reactive oxygen species (ROS) production, as well as changes in the physiology and the characteristics of the algal cells, including mortality, size, internal complexity and chlorophyll fluorescence. These experiments suggest different effects of the several species of harmful algae upon bivalve haemocytes. Some harmful algae act as immunostimulants, whereas others are immunosuppressive. P. minimum appears to activate haemocytes, but the other harmful algal species tested seem to cause a suppression of immune functions, generally consisting of decreases in phagocytosis, production of ROS and cell adhesion and besides cause an increase in the percentage of dead haemocytes, which could be attributable to the action of chemical toxins. Microalgal cells exposed to shellfish haemolymph generally showed evidence of algal degradation, e.g. loss of chlorophyll fluorescence and modification of cell shape. Thus, in vitro tests allow a better understanding of the role of the haemocytes and the haemolymph in the defence mechanisms protecting molluscan shellfish from harmful algal cells and could also be further developed to estimate the effects of HABs on bivalve molluscs in vivo.     

Resolving the complex relationship between harmful algal blooms and environmental factors in the coastal waters adjacent to the Changjiang River estuary

The sea area adjacent to the Changjiang River estuary is the most notable region for harmful algal blooms (HABs¹) in China as both diatom and dinoflagellate blooms have been recorded in this region. Affected by the Changjiang diluted water (CDW²) and currents from the open ocean (i.e., Taiwan warm current, TWC³), the environmental conditions in the coastal waters adjacent to the Changjiang River estuary are quite complex. To obtain a better understanding of the mechanisms of HABs in this region, analyses based on field investigation data collected by the National Basic Research Priority Program (CEOHAB I⁴) were performed using principle component analysis (PCA⁵), multiple regression analysis (MRA⁶) and path analysis (PA⁷). The results suggested that phosphate and silicate are the major factors that directly affect the diatom bloom, while dissolved inorganic nitrogen (DIN⁸), temperature and turbidity are the factors that influence the dinoflagellate bloom. CDW and the TWC have different roles in affecting the two types of algal blooms. CDW, which has a high concentration of nitrate and silicate, is essential for the diatom bloom, while the intrusion of the TWC (mainly Kuroshio subsurface water that is rich in phosphate at the bottom) is critical for the maintenance of the dinoflagellate bloom. The results of this study offer a better understanding of the mechanisms of HABs in the East China Sea.     

Detection of bloom-forming dinoflagellates Karenia mikimotoi and Prorocentrum donghaiense using qPCR assays

Journal of Applied Phycology - The dinoflagellates Karenia mikimotoi and Prorocentrum donghaiense are both important causative species of harmful algal blooms (HABs) in the East China Sea. The...     

Satellite discrimination of Karenia mikimotoi and Phaeocystis harmful algal blooms in European coastal waters: Merged classification of ocean colour data

The detection of dense harmful algal blooms (HABs) by satellite remote sensing is usually based on analysis of chlorophyll-a as a proxy. However, this approach does not provide information about the potential harm of bloom, nor can it identify the dominant species. The developed HAB risk classification method employs a fully automatic data-driven approach to identify key characteristics of water leaving radiances and derived quantities, and to classify pixels into “harmful”, “non-harmful” and “no bloom” categories using Linear Discriminant Analysis (LDA). Discrimination accuracy is increased through the use of spectral ratios of water leaving radiances, absorption and backscattering. To reduce the false alarm rate the data that cannot be reliably classified are automatically labelled as “unknown”. This method can be trained on different HAB species or extended to new sensors and then applied to generate independent HAB risk maps; these can be fused with other sensors to fill gaps or improve spatial or temporal resolution. The HAB discrimination technique has obtained accurate results on MODIS and MERIS data, correctly identifying 89% of Phaeocystis globosa HABs in the southern North Sea and 88% of Karenia mikimotoi blooms in the Western English Channel. A linear transformation of the ocean colour discriminants is used to estimate harmful cell counts, demonstrating greater accuracy than if based on chlorophyll-a; this will facilitate its integration into a HAB early warning system operating in the southern North Sea.     

Sexual resting cyst production by the dinoflagellate Akashiwo sanguinea: a potential mechanism contributing to the ubiquitous distribution of a harmful alga

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.     

Severe Karenia brevis red tides influence juvenile bottlenose dolphin (Tursiops truncatus) behavior in Sarasota Bay,Florida

Harmful algal blooms (HABs) are natural stressors in the coastal environment that may be increasing in frequency and severity. This study investigates whether severe red tide blooms, caused by Karenia brevis, affect the behavior of resident coastal bottlenose dolphins in Sarasota Bay, Florida through changes to juvenile dolphin activity budgets, ranging patterns, and social associations. Behavioral observations were conducted on free‐ranging juvenile dolphins during the summer months of 2005–2007, and behavior during red tide blooms was compared to periods of background K. brevis abundance. We also utilized dolphin group sighting data from 2004 to 2007 to obtain comparison information from before the most severe recent red tide of 2005 and incorporate social association information from adults in the study area. We found that coastal dolphins displayed a suite of behavioral changes associated with red tide blooms, including significantly altered activity budgets, increased sociality, and expanded ranging behavior. At present, we do not fully understand the mechanism behind these red tide‐associated behavioral effects, but they are most likely linked to underlying changes in resource availability and distribution. These behavioral changes have implications for more widespread population impacts, including increased susceptibility to disease outbreaks, which may contribute to unusual mortality events during HABs.     

The Diversity of Harmful Algal Bloom-Triggering Mechanisms and the Complexity of Bloom Initiation

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.     

Vulnerability of coastal ecosystems to changes in harmful algal bloom distribution in response to climate change: projections based on model analysis

Harmful algal blooms (HABs), those proliferations of algae that can cause fish kills, contaminate seafood with toxins, form unsightly scums, or detrimentally alter ecosystem function have been increasing in frequency, magnitude, and duration worldwide. Here, using a global modeling approach, we show, for three regions of the globe, the potential effects of nutrient loading and climate change for two HAB genera, pelagic Prorocentrum and Karenia, each with differing physiological characteristics for growth. The projections (end of century, 2090–2100) are based on climate change resulting from the A1B scenario of the Intergovernmental Panel on Climate Change Institut Pierre Simon Laplace Climate Model (IPCC, IPSL‐CM4), applied in a coupled oceanographic‐biogeochemical model, combined with a suite of assumed physiological ‘rules’ for genera‐specific bloom development. Based on these models, an expansion in area and/or number of months annually conducive to development of these HABs along the NW European Shelf‐Baltic Sea system and NE Asia was projected for both HAB genera, but no expansion (Prorocentrum spp.), or actual contraction in area and months conducive for blooms (Karenia spp.), was projected in the SE Asian domain. The implications of these projections, especially for Northern Europe, are shifts in vulnerability of coastal systems to HAB events, increased regional HAB impacts to aquaculture, increased risks to human health and ecosystems, and economic consequences of these events due to losses to fisheries and ecosystem services.     

Applications of Satellite Ocean Color Sensors for Monitoring and Predicting Harmful Algal Blooms

The new satellite ocean color sensors offer a means of detecting and monitoring algal blooms in the ocean and coastal zone. Beginning with SeaWiFS (Sea Wide Field-of-view Sensor) in September 1997, these sensors provide coverage every 1 to 2 days with 1-km pixel view at nadir. Atmospheric correction algorithms designed for the coastal zone combined with regional chlorophyll algorithms can provide good and reproducible estimates of chlorophyll, providing the means of monitoring various algal blooms. Harmful algal blooms (HABs) caused by Karenia brevis in the Gulf of Mexico are particularly amenable to remote observation. The Gulf of Mexico has relatively clear water and K. brevis, in bloom conditions, tends to produce a major portion of the phytoplankton biomass. A monitoring program has begun in the Gulf of Mexico that integrates field data from state monitoring programs with satellite imagery, providing an improved capability for the monitoring of K. brevis blooms.     

Noctiluca scintillans may act as a vector of toxigenic microalgae

Noctiluca scintillans food vacuoles containing toxigenic microalgae of the genera Dinophysis and Pseudo-nitzschia are reported in samples from the Galician Rías Baixas (NW Spain). N. scintillans may play an important role in the population dynamics of toxigenic microalgae blooms, and may act as a vector of phycotoxins to higher trophic levels. The harmful effects of Noctiluca in natural shellfish beds and/or aquaculture sites may be magnified if cells are loaded with toxigenic microalgal prey.     

High Resolution Spatio-temporal Detection of Potentially Harmful Dinoflagellates in Confined Waters of the NW Mediterranean

A systematic sampling programme was carried out in a large numberof confined waters (principally harbours) along the Catalancoast (NW Mediterranean) in the context of a new MonitoringProgramme. This Monitoring Programme was associated not onlywith areas subject to aquaculture activities, and thereforeunder legislation, but also with confined areas with a highrisk of harmful algal blooms (HABs) occurrence, in order toprovide an early warning of potentially widespread HABs. Thesystematic programme was performed weekly in summer and bi-monthlyin winter for five years. The main results were: (i) the detectionof many harmful species and the presence of high numbers ofharmful dinoflagellates, mainly of the genera Alexandrium andDinophysis; (ii) the detection of Alexandrium catenella, newin the study area, which had hardly ever been detected in theMediterranean Sea; (iii) the presence of some potentially harmfulspecies, including Dinophysis sacculus, present at all periodsof the year; (iv) bloom recurrence in several stations; (v)occasional coincidence of small-scale blooms, such as thoseconfined inside the harbours, with widespread blooms (mesoscaleblooms) of the same organism. The implications of this highfrequency of HAB detection is discussed in relation to the suitabilityof this sampling programme (focused on confined waters) forthe early detection of algal blooms.     

Zooming on dynamics of marine microbial communities in the phycosphere of Akashiwo sanguinea (Dinophyta) blooms

Characterizing ecological relationships between viruses, bacteria and phytoplankton in the ocean is critical to understanding the ecosystem; however, these relationships are infrequently investigated together. To understand the dynamics of microbial communities and environmental factors in harmful algal blooms (HABs), we examined the environmental factors and microbial communities during Akashiwo sanguinea HABs in the Jangmok coastal waters of South Korea by metagenomics. Specific bacterial species showed complex synergistic and antagonistic relationships with the A. sanguinea bloom. The endoparasitic dinoflagellate Amoebophrya sp. 1 controlled the bloom dynamics and correlated with HAB decline. Among nucleocytoplasmic large DNA viruses (NCLDVs), two Pandoraviruses and six Phycodnaviruses were strongly and positively correlated with the HABs. Operational taxonomic units of microbial communities and environmental factors associated with A. sanguinea were visualized by network analysis: A. sanguinea–Amoebophrya sp. 1 (r = .59, time lag: 2 days) and A. sanguinea–Ectocarpus siliculosus virus 1 in Phycodnaviridae (0.50, 4 days) relationships showed close associations. The relationship between A. sanguinea and dissolved inorganic phosphorus relationship also showed a very close correlation (0.74, 0 day). Microbial communities and the environment changed dynamically during the A. sanguinea bloom, and the rapid turnover of microorganisms responded to ecological interactions. A. sanguinea bloom dramatically changes the environments by exuding dissolved carbohydrates via autotrophic processes, followed by changes in microbial communities involving host‐specific viruses, bacteria and parasitoids. Thus, the microbial communities in HAB are composed of various organisms that interact in a complex manner.     

沿岸海域富营养化与赤潮发生的关系

综述了赤潮的发生与沿岸海域富营养化的关系。近几十年来,人类活动使得天然水体的富营养化进程大大加速。营养负荷的增加与高生物量水华的增多相联系。控制营养输入后,浮游植物生物量或有害藻类水华事件也相应减少。营养的组成与浮游植物的种类组成及水华的形成有密切联系。有机营养对有害藻类水华的促进作用受到关注。营养输入时机影响浮游植物种间竞争的结果,因而对浮游植物的群落演替具有深远影响。由于浮游植物存在生理差异,因而对营养加富的反应因种而异。营养在调控某些有毒藻类的毒素产量方面也发挥着重要作用。此外,营养输入与藻类水华之间存在复杂的间接联系。当然,营养状况并非浮游植物群落演替的唯一决定因素。研究结果提示,控制营养输入、减缓水域富营养化是减少有害藻类水华发生的有效途径,而深入研究典型有害藻类的营养生理对策则为防治并最终消除有害藻类水华提供了理论基础。     

Fish Sound Production in the Presence of Harmful Algal Blooms in the Eastern Gulf of Mexico

This paper presents the first known research to examine sound production by fishes during harmful algal blooms (HABs). Most fish sound production is species-specific and repetitive, enabling passive acoustic monitoring to identify the distribution and behavior of soniferous species. Autonomous gliders that collect passive acoustic data and environmental data concurrently can be used to establish the oceanographic conditions surrounding sound-producing organisms. Three passive acoustic glider missions were conducted off west-central Florida in October 2011, and September and October 2012. The deployment period for two missions was dictated by the presence of red tide events with the glider path specifically set to encounter toxic Karenia brevis blooms (a.k.a red tides). Oceanographic conditions measured by the glider were significantly correlated to the variation in sounds from six known or suspected species of fish across the three missions with depth consistently being the most significant factor. At the time and space scales of this study, there was no detectable effect of red tide on sound production. Sounds were still recorded within red tide-affected waters from species with overlapping depth ranges. These results suggest that the fishes studied here did not alter their sound production nor migrate out of red tide-affected areas. Although these results are preliminary because of the limited measurements, the data and methods presented here provide a proof of principle and could serve as protocol for future studies on the effects of algal blooms on the behavior of soniferous fishes. To fully capture the effects of episodic events, we suggest that stationary or vertically profiling acoustic recorders and environmental sampling be used as a complement to glider measurements.     

Detection of several harmful algal species by sandwich hybridization integrated with a nuclease protection assay

The frequent occurrence of harmful algal blooms (HABs) is a pressing topic in marine research. An integrated sandwich hybridization and nuclease protection assay was established to qualitatively and quantitatively detect 12 harmful algal species. This method demonstrated good reliability, specificity and accuracy for analyzing samples from individual and mixed cultures, as well as field collection, and cell volumes were positively correlated to the slopes of calibration curves. The lowest quantitative detection limits were those concentrations observed during blooms; thus, this technique provides an efficient alternative to microscopy for rapid identification and quantitation of harmful algal species and could be routinely used to monitor phytoplankton in field surveys.     

东海原甲藻与中肋骨条藻的种间竞争数值模拟

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

Recombinase-aided amplification combined with lateral flow dipstick for the rapid detection of Amphidinium carterae

Journal of Applied Phycology - In the context of global climate change, the frequency and duration of harmful algal blooms (HABs) due to eutrophic coastal waters have increased. HABs can cause...     

Eutrophication and harmful algal blooms: A scientific consensus

In January 2003, the US Environmental Protection Agency sponsored a “roundtable discussion” to develop a consensus on the relationship between eutrophication and harmful algal blooms (HABs), specifically targeting those relationships for which management actions may be appropriate. Academic, federal, and state agency representatives were in attendance. The following seven statements were unanimously adopted by attendees based on review and analysis of current as well as pertinent previous data:

• (1)Degraded water quality from increased nutrient pollution promotes the development and persistence of many HABs and is one of the reasons for their expansion in the U.S. and other nations;
• (2)The composition–not just the total quantity–of the nutrient pool impacts HABs;
• (3)High-biomass blooms must have exogenous nutrients to be sustained;
• (4)Both chronic and episodic nutrient delivery promote HAB development;
• (5)Recently developed tools and techniques are already improving the detection of some HABs, and emerging technologies are rapidly advancing toward operational status for the prediction of HABs and their toxins;
• (6)Experimental studies are critical to further the understanding about the role of nutrients in HABs expression, and will strengthen prediction and mitigation of HABs; and
• (7)Management of nutrient inputs to the watershed can lead to significant reduction in HABs.

Supporting evidence and pertinent examples for each consensus statement are provided herein.