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.     

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.     

具齿原甲藻的生态特征及赤潮成因浅析

在东海的长江口和舟山海域,2000-2002年连续3年的5月发生罕见的大规模赤潮,赤潮生物为具齿原甲藻,为了解该种的生态特征及赤潮频发原因,本文分析近赤潮应急监测资料及野外生态调查资料,结果表明,该种适温,适盐范围分别为15-25℃和14-32,最适温度,盐度分别为18-22℃和22-31,该种运动能力较强,在海流交汇的紊乱水体中具有强适应能力,具有昼夜垂直移动特性,水温,硝酸盐含量,海流及上升流,磷限制环境下的种间竞争优势对诱发和维持赤潮具有重要作用.     

南海台风引发藻华的生物机制

海洋浮游植物的生长和小型浮游动物的摄食,与温度、光照和营养盐等因素密切相关。中国南海海盆是层化结构稳定的寡营养海,然而每年台风过后都有大量藻华的报道。为了探究台风引发藻华的生物形成机制,通过南海海域表层和次表层叶绿素最大层(DCM)中的小型浮游动物的摄食活动以及浮游植物对光照和营养响应的现场实验,探究小型浮游动物摄食、光照和营养对海洋浮游植物生长和群落结构的调控作用。实验表明:1)表层水体中的小型浮游动物摄食速率明显大于DCM层;小型浮游动物摄食在表层水体中以大粒级的浮游植物(5μm)为主,在DCM层中没有明显粒级选择;自然光中的紫外线能增加小型浮游动物的摄食;营养盐添加轻微降低小型浮游动物的摄食。2)营养盐和光照的增强显著促进DCM层浮游植物的生长,并增加大粒级浮游植物的占比;而光照和营养盐变化对表层浮游植物的生长和粒级结构的改变相对不明显。因此,相对表层,深层浮游植物面临较小的摄食压力,拥有更大的光照和营养盐需求潜能;当台风引发水体垂向混合后,获得营养盐补充的表层浮游植物并不能迅速生长,而获得充足光照的深层浮游植物能迅速生长,成为藻华爆发的优势种。     

Dinoflagellate cysts from surface sediments of Saldanha Bay, South Africa: an indication of the potential risk of harmful algal blooms

The distribution and abundance of dinoflagellate cysts from recent coastal sediments in Saldanha Bay, was investigated, and compared to the cyst assemblages of the adjacent coastal upwelling system as reflected in the sediments off Lambert's Bay on the southern Namaqua shelf. Twenty-two cyst types were identified from three sample sites off Lambert's Bay with recorded abundances between 1726 and 1863 cysts ml⁻¹ wet sediment. At least 21 distinctive cyst types were identified from 32 sample sites within Saldanha Bay. Cyst abundance in Saldanha Bay was relatively low, averaging 116 cysts ml⁻¹ wet sediment. The region off Lambert's Bay is especially susceptible to the formation of harmful algal blooms attributed to high biomass dinoflagellate blooms. Owing to these blooms and the retentive circulation characteristics of this area, cyst formation and deposition is high. Blooms can be advected into Saldanha Bay, but their development and duration in the Bay is restricted by the system of exchange that operates between the Bay and the coastal upwelling system, in that there is a net export of surface waters from the Bay. Consequently, fewer cysts are formed and deposited within the Bay thereby reducing the likelihood of in situ bloom development initiated from the excystment of cysts.     

The value of harmful algal bloom predictions to the nearshore commercial shellfish fishery in the Gulf of Maine

In this study, we develop a framework for measuring the value of harmful algal bloom (HAB) predictions. The framework captures the effects of both private and public responses to HABs. Using data from the New England nearshore commercial shellfish fishery and impact estimates for a large-scale HAB event in 2005, we illustrate how the potential value of HAB forecasts may be estimated. The results of our study suggest that the long-term value of a HAB prediction and tracking system for the Gulf of Maine is sensitive to the frequency of HAB events, the accuracy of predictions, the choice of HAB impact measures, and the effectiveness of public and private responses.     

A model for the prediction of harmful algae blooms in the Vietnamese upwelling area

Satellite pictures and in situ observations indicate strong phytoplankton blooms including harmful algae blooms (HABs) during southwest (SW) summer monsoon in the Vietnamese upwelling area. In this period, nutrients are provided by coastal upwelling and by the very high river runoff from the Mekong River. During SW monsoon, in general two circulation patterns exist which allow the prediction of advection and diffusion of HAB patches. A Lagrangian HAB model that is driven by a circulation model and applied to HABs in Vietnamese waters is presented. Advection which is the most complicated part in modelling transport of passive substances is validated with a Lagrangian sediment trap experiment. The model produces realistic results compared to in situ observations and satellite images and might be used for real time forecast in the future.     

大亚湾春季卡盾藻种群动态及其赤潮成因分析

研究了2000年春季大亚湾古老卡盾藻/海洋卡盾藻(Chattonella antique/marina)种群动态及其与环境因子的关系。卡盾藻是一种沿岸分布的有害赤潮针孢藻,2000年春在大亚湾海域高密度高频率出现,最高细胞密度为129cells/mL。大亚湾合适的水温、盐度及丰富的营养盐含量为卡盾藻的生长提供了良好的环境条件,而较低的Si∶N及持续的阴雨天气抑制了小型硅藻的生长,使卡盾藻在浮游植物种群竞争中占据一定地位。孢囊萌发被认为是水体中卡盾藻营养细胞及其赤潮发生的种源,但孢囊需经过4个月以上、11℃以下低温处理后方可成熟和萌发,大亚湾冬季水温,不能满足卡盾藻孢囊休眠成熟的需要。作者推测,2000年春大亚湾高密度卡盾藻营养细胞的种源可能来源于由船舶压舱水所带来的成熟孢囊。       

山东海域赤潮灾害特征及预警报管理

掌握赤潮灾害时空分布规律是科学设计赤潮监控方案、提高防灾管理效率的基础和依据。本文统计分析了1990—2010年间山东海域的76次赤潮灾害事件数据。结果表明:赤潮灾害频率显著地分为4个波段,且峰值呈递增趋势;灾害面积呈现周期性倒U型曲线变化;每年的5—10月是赤潮灾害的多发期,其中,8月的灾害次数和灾害面积均占全年总数的30%以上;灾害空间分布集中在莱州湾海域、青岛近海、庙岛群岛的北隍城海域,烟台四十里湾的赤潮灾害最频繁;夜光藻(Noctiluca scintillans)和球型棕囊藻(Phaeocystis globosa)引发的赤潮灾害面积最高,夜光藻和红色裸甲藻(Gymnodinium sanguineum)致灾次数最多。本文从赤潮监测与信息采集、赤潮预报方法以及赤潮灾害预警报管理机制3个方面分析总结了山东海域赤潮灾害预警报业务化管理现状,结合山东海域赤潮灾害特征和预警报管理实践,提出了以控制污染和修复生态环境为根本的赤潮防控建议。     

Genomewide investigation of adaptation to harmful algal blooms in common bottlenose dolphins (Tursiops truncatus)

Harmful algal blooms (HABs), which can be lethal in marine species and cause illness in humans, are increasing worldwide. In the Gulf of Mexico, HABs of Karenia brevis produce neurotoxic brevetoxins that cause large‐scale marine mortality events. The long history of such blooms, combined with the potentially severe effects of exposure, may have produced a strong selective pressure for evolved resistance. Advances in next‐generation sequencing, in particular genotyping‐by‐sequencing, greatly enable the genomic study of such adaptation in natural populations. We used restriction site‐associated DNA (RAD) sequencing to investigate brevetoxicosis resistance in common bottlenose dolphins (Tursiops truncatus). To improve our understanding of the epidemiology and aetiology of brevetoxicosis and the potential for evolved resistance in an upper trophic level predator, we sequenced pools of genomic DNA from dolphins sampled from both coastal and estuarine populations in Florida and during multiple HAB‐associated mortality events. We sequenced 129 594 RAD loci and analysed 7431 single nucleotide polymorphisms (SNPs). The allele frequencies of many of these polymorphic loci differed significantly between live and dead dolphins. Some loci associated with survival showed patterns suggesting a common genetic‐based mechanism of resistance to brevetoxins in bottlenose dolphins along the Gulf coast of Florida, but others suggested regionally specific mechanisms of resistance or reflected differences among HABs. We identified candidate genes that may be the evolutionary target for brevetoxin resistance by searching the dolphin genome for genes adjacent to survival‐associated SNPs.     

Harmful algal bloom (HAB) in the East Sea identified by the Geostationary Ocean Color Imager (GOCI)

Harmful Cochlodinium polykrikoides blooms have frequently appeared and caused fatal harm to aquaculture in Korean coastal waters since 1995. We investigated the applicability of GOCI, the world's first Geostationary Ocean Color Imager, in monitoring the distribution and temporal movement of a harmful algal bloom (HAB) that was discovered in the East Sea near the Korean peninsula in August 2013. We identified the existence of C. polykrikoides at a maximum cell abundance of over 6000 cells/mL and a chlorophyll a concentration of over 400 mg/m³. In areas of C. polykrikoides blooms, GOCI remote sensing reflectance (R_rs) spectra demonstrated the typical radiometric features of a HAB, and from the diurnal variations using GOCI-derived chlorophyll concentration images, we were able to identify the vertical migration of the red tide species. We also found that the formation and propagation of the HAB had relations with cold water mass in the coastal region. GOCI can be effectively applied to the monitoring of short-term and long-term movements of red tides.     

东海赤潮高发区沉积物中营养盐再生速率的研究

2001年5月和2002年5月在东海赤潮高发区的4个站位(E4、E5、DB6、DCl0)对沉积物表面分别充氮气和空气进行培养,研究沉积物．水界面营养盐交换通量在不同条件下的变化规律,以及该海域沉积物中营养盐的再生对水体中营养盐的贡献．结果表明,各溶解态营养盐在还原条件下的迁移较为活跃．在距离陆地较近的海域,营养盐一般由水向沉积物中扩散,且距陆地越近,交换通量越大;而在上升流区,营养盐多由沉积物中向水中扩散．东海赤潮高发区沉积物是SiO3^2-的源,对初级生产力的贡献占6％．同时,东海赤潮高发区沉积物是氮、磷营养盐蓄积库．该海域沉积物每年从水体中吸附的DIN、PO4^3-分别占长江输入的5．9％、67％,沉积物一水界面对水体中SiO3^2-的贡献占7．8％．       

Potentially toxic and harmful microalgae from coastal waters of the Campania region (Tyrrhenian Sea, Mediterranean Sea)

A total number of 40 potentially toxic microalgae and 5 taxa causing discolorations have been identified along the coasts of the Campania region (South Tyrrhenian Sea, Mediterranean Sea). This number is based on results of over 20 years of research at a coastal station in the Gulf of Naples, 4 years of monitoring activity along the Campania coasts and of a series of scattered and sporadic observations. Several species of the recently erected genera Karenia and Takayama are reported for the first time in the area. Information on the period of highest abundances or most probable period of occurrence indicates the late spring and summer as the periods of maximum risk of harmful events. Despite the variety of potentially toxic species, no human health problems nor fish kills have ever been recorded in Campania. The reasons for this apparent paradox are probably to be found in the ecological factors that regulate the abundance, toxicity and spatial distribution of the potentially harmful species and reflect as well the relatively low number of aquaculture plants in the area.     

Reflections on the Interpretation and Application of Article 121(3) in the South China Sea Arbitration (Merits)

The 2016 South China Sea Arbitration Award provided a detailed review of the interpretation of Article 121(3) of the U.N. Convention on the Law of the Sea. According to the Tribunal, Article 121(3) performs a preventive function by disabling tiny features from unfairly generating enormous entitlements to maritime space that does not serve the local population. This provision also contributes to safeguarding of the Common Heritage of Mankind. The Tribunal's interpretation seems to reflect development of the law of the sea toward protection of the common interests of the international community. However, since the Tribunal's interpretation is not anchored in state practice and the jurisprudence, whether the interpretation can be generalized needs careful consideration.     

一株南海指海绵共附生真菌多样性及其抗菌活性

本研究通过采用6种不同的培养基对南海指海绵上的共附生真菌进行平板涂布法分离培养,结合ITS-rDNA基因序列特征和形态学特征进行鉴定来研究海绵共附生真菌多样性。共分离培养出55株(21种)真菌菌株,优势菌属为木霉菌属Trichoderma sp.(31株),其中的哈茨木霉Trichoderma harzianum 22株作为优势菌种,占真菌菌株总数的40%。其余真菌则属于枝孢菌属Cladosporium、迈耶氏酵母属Meyerozyma、茎点霉属Phoma、红酵母属Rhodotorula、青霉菌属Penicillium、曲霉属Aspergillus等10个属,还有两株未鉴定到种的子囊菌。选取金黄色葡萄球菌Staphylococcus aureus等7种病原指示菌,采用琼脂扩散法对21种真菌的代表性菌株进行抗菌活性筛选,其中有3种真菌至少对一种指示菌表现出抑菌效果,占菌株种类总数的11%。     

中国南海沉积环境可培养细菌多样性研究

【目的】探索海洋沉积环境中可培养细菌的多样性。【方法】采用纯培养分离及16S rRNA基因序列鉴定的方法,对我国南海海域20个沉积物样品进行细菌多样性分析。【结果】共获得200株细菌,分属于47个属,99个种。经系统进化分析,可培养菌株主要分布于4个类群:厚壁菌门(Firmicutes)、变形菌门(Proteobacteria)、放线菌门(Actinobacteria)和拟杆菌门(Bacteroidetes),优势类群为Firmicutes,其中芽孢杆菌属(Bacillus)所占比例为55.6%;而Actinobacteria和Bacteroidetes两个类群获得菌株较少;在Firmicutes和Actinobacteria两个类群中发现8个潜在新种和3个潜在新属级类群。【结论】初步研究结果表明,南海海洋沉积环境可培养微生物资源丰富,新物种资源多样;其中,芽孢杆菌为海洋沉积环境中的优势类群,随着样品深度的增加,细菌多样性呈现递减的趋势,深度可能是影响细菌多样性的一个重要因素;其次,分离培养基和分离方法直接关系到样品中可培养微生物多样性的发现,有待深入研究。     

The potential threat of algal blooms to the abalone (Haliotis midae) mariculture industry situated around the South African coast

Toxic algal blooms are common world-wide and pose a serious problem to the aquaculture and fishing industries. Dinoflagellate species such as Karenia brevis, Karenia mikimotoi, Heterosigma akashiwo and Chatonella cf. antiqua are recognised toxic species implicated in various faunal mortalities. Toxic blooms of Karenia cristata were observed on the south coast of South Africa for the first time in 1988 and were responsible for mortalities of wild and farmed abalone. K. cristata and various other dinoflagellate species common along the South African coast, as well as K. mikimotoi (Isolation site: Norway, Univ. of Copenhagen) and K. brevis (Isolation site: Florida, BIGELOW), were tested for toxicity by means of a bioassay involving Artemia larvae as well as abalone larvae and spat. K. cristata, like K. brevis, contains an aerosol toxin; however, the toxin present in K. cristata has not yet been isolated and remains unknown. K. brevis was, therefore, used to determine which developmental phase of the bloom would affect abalone farms most, and whether ozone could be used as an effective mitigating agent. Of the 17 dinoflagellate species tested, K. cristata, Akashiwo sanguinea, K. mikimotoi and K. brevis pose the greatest threat to the abalone mariculture industry. K. brevis was most toxic during its exponential and stationary phases. Results suggest that ozone is an effective mitigation agent but its economic viability for use on abalone farms must still be investigated.     

Mixotrophy in the phototrophic dinoflagellate Takayama helix (family Kareniaceae): Predator of diverse toxic and harmful dinoflagellates

Takayama spp. are phototrophic dinoflagellates belonging to the family Kareniaceae and have caused fish kills in several countries. Understanding their trophic mode and interactions with co-occurring phytoplankton species are critical steps in comprehending their ecological roles in marine ecosystems, bloom dynamics, and dinoflagellate evolution. To investigate the trophic mode and interactions of Takayama spp., the ability of Takayama helix to feed on diverse algal species was examined, and the mechanisms of prey ingestion were determined. Furthermore, growth and ingestion rates of T. helix feeding on the dinoflagellates Alexandrium lusitanicum and Alexandrium tamarense, which are two optimal prey items, were determined as a function of prey concentration. T. helix ingested large dinoflagellates ≥15 μm in size, except for the dinoflagellates Karenia mikimotoi, Akashiwo sanguinea, and Prorocentrum micans (i.e., it fed on Alexandrium minutum, A. lusitanicum, A. tamarense, A. pacificum, A. insuetum, Cochlodinium polykrikoides, Coolia canariensis, Coolia malayensis, Gambierdiscus caribaeus, Gymnodinium aureolum, Gymnodinium catenatum, Gymnodinium instriatum, Heterocapsa triquetra, Lingulodinium polyedrum, and Scrippsiella trochoidea). All these edible prey items are dinoflagellates that have diverse eco-physiology such as toxic and non-toxic, single and chain forming, and planktonic and benthic forms. However, T. helix did not feed on small flagellates and dinoflagellates <13 μm in size (i.e., the prymnesiophyte Isochrysis galbana; the cryptophytes Teleaulax sp., Storeatula major, and Rhodomonas salina; the raphidophyte Heterosigma akashiwo; the dinoflagellates Heterocapsa rotundata, Amphidinium carterae, Prorocentrum minimum; or the small diatom Skeletonema costatum). T. helix ingested Heterocapsa triquetra by direct engulfment, but sucked materials from the rest of the edible prey species through the intercingular region of the sulcus. With increasing mean prey concentration, the specific growth rates of T. helix on A. lusitanicum and A. tamarense increased continuously before saturating at prey concentrations of 336–620 ng C mL⁻¹. The maximum specific growth rates (mixotrophic growth) of T. helix on A. lusitanicum and A. tamarense were 0.272 and 0.268 d⁻¹, respectively, at 20 °C under a 14:10 h light/dark cycle of 20 μE m⁻² s⁻¹ illumination, while its growth rates (phototrophic growth) under the same light conditions without added prey were 0.152 and 0.094 d⁻¹, respectively. The maximum ingestion rates of T. helix on A. lusitanicum and A. tamarense were 1.23 and 0.48 ng C predator⁻¹d⁻¹, respectively. The results of the present study suggest that T. helix is a mixotrophic dinoflagellate that is able to feed on a diverse range of toxic species and, thus, its mixotrophic ability should be considered when studying red tide dynamics, food webs, and dinoflagellate evolution.     

Population genetic structure and connectivity of the harmful dinoflagellate Alexandrium minutum in the Mediterranean Sea

The toxin-producing microbial species Alexandrium minutum has a wide distribution in the Mediterranean Sea and causes high biomass blooms with consequences on the environment, human health and coastal-related economic activities. Comprehension of algal genetic differences and associated connectivity is fundamental to understand the geographical scale of adaptation and dispersal pathways of harmful microalgal species. In the present study, we combine A. minutum population genetic analyses based on microsatellites with indirect connectivity (C(i)) estimations derived from a general circulation model of the Mediterranean sea. Our results show that four major clusters of genetically homogeneous groups can be identified, loosely corresponding to four regional seas: Adriatic, Ionian, Tyrrhenian and Catalan. Each of the four clusters included a small fraction of mixed and allochthonous genotypes from other Mediterranean areas, but the assignment to one of the four clusters was sufficiently robust as proved by the high ancestry coefficient values displayed by most of the individuals (>84%). The population structure of A. minutum on this scale can be explained by microalgal dispersion following the main regional circulation patterns over successive generations. We hypothesize that limited connectivity among the A. minutum populations results in low gene flow but not in the erosion of variability within the population, as indicated by the high gene diversity values. This study represents a first and new integrated approach, combining both genetic and numerical methods, to characterize and interpret the population structure of a toxic microalgal species. This approach of characterizing genetic population structure and connectivity at a regional scale holds promise for the control and management of the harmful algal bloom events in the Mediterranean Sea.     

大亚湾海域锥状斯氏藻孢囊形成与萌发的季节变化

锥状斯氏藻(Scrippsiella trochoidea)是南海大亚湾海域优势甲藻。为了解该藻孢囊形成和萌发动态及其对营养细胞种群动态的影响,2001年1月-2002年1月在大亚湾澳头海域用沉积物捕捉器及TFO重力采泥器对其孢囊进行每月一次的周年监测,同时对浮游植物、水温、盐度、溶解氧等也进行了监测。孢囊形成和萌发分别以沉积物捕捉器中的孢囊形成率以及上表层沉积物中空孢囊的百分比来表示。钙质孢囊和非钙质孢囊年平均形成率分别为1.11×104 cysts m-2d-1和2.13×105 cysts m-2d-1。前者在冬季大量形成,而后者在夏季形成较多。孢囊多在春秋季节萌发,夏季萌发较少,而冬季几乎不萌发。在5月份和10月份营养细胞数量峰形成前,孢囊的萌发出现了高峰,而表层沉积物中的孢囊数量及孢囊形成率则在营养细胞数量峰后大幅度上升。由此可见,大亚湾沉积物中该藻孢囊的萌发给水体提供了丰富的营养细胞,反之水体中高密度营养细胞又促使孢囊的大量形成,从而造成了锥状斯氏藻赤潮在大亚湾海域接连发生。