赤潮影响因素研究进展

通过对赤潮发生的生物、化学、物理等影响因素研究现状的分析,认为赤潮生物是引发赤潮的内在因素,化学因素是赤潮发生的物质基础,而物理因素则是提供赤潮发生的环境条件。在赤潮生物存在的前提下,化学和物理因素都可以限制赤潮的发生,所以,将生物学、化学与物理海洋学耦合仍旧是赤潮研究需要解决的问题。在分析了现有研究状况的基础上,从不同角度提出了赤潮的研究方向。     

有害藻华的宏条形码分析:机会与挑战

宏条形码分析(metabarcoding analysis)是通过对野外样本的分子标记(即条形码)进行PCR扩增、测序,并利用测序结果分析野外样本物种组成、物种相对丰度,及其时空动态变化。这项在近20年里逐步发展起来的分子技术,得益于DNA测序技术的快速发展,包括测序通量的提升和测序成本的降低以及生物信息学分析方法的快速开发。基于可操作分类单元(operational taxonomic units,OTUs)分析方法,特别是最近的扩增子序列变异(amplicon sequence variants,ASVs)方法的开发,极大促进了宏条形码分析的应用。在世界及我国的海洋生态领域,宏条形码分析的应用还处于起步阶段。随着分析方法的日趋成熟,可适用于不同应用领域的分子标记不断增加,以及参考数据库的不断完善,宏条形码分析将释放出广阔的应用潜力。可以预期,宏条形码分析将在系统分析我国近岸海域有害藻华物种(及其他浮游生物物种)的组成和相对丰度中起关键作用,通过发现新物种,辨别隐存种,跟踪物种的生物地理学,进而帮助解析有害藻华暴发机理。     

Discussion about mechanism of harmful algal blooms breakout

HAB (harmful algal bloom) is a serious marine ecological disaster. Up to now there is no definite conclusion about its mechanism of occurrence.The observation results show that the HAB breakout in the Xiangshan Bay was mainly caused by physical convergence capacity,and the breakout process had no direct relation to eutrophication.As a new idea it is thought that the process of the HAB breakout is mainly a physical convergence or accumulation process in some areas. A hypothesis about dynamic mechanism of the HAB appearing in the area off the Changjiang Estuary is put forward according to hydrology and topography and the past work, and a breakthrough is expected to be made for doing further research.     

Approaches to monitoring,control and management of harmful algal blooms (HABs)

Virtually every coastal country in the world is affected by harmful algal blooms (HABs, commonly called “red tides”). These phenomena are caused by blooms of microscopic algae. Some of these algae are toxic, and can lead to illness and death in humans, fish, seabirds, marine mammals, and other oceanic life, typically as a result of the transfer of toxins through the food web. Sometimes the direct release of toxic compounds can be lethal to marine animals. Non-toxic HABs cause damage to ecosystems, fisheries resources, and recreational facilities, often due to the sheer biomass of the accumulated algae. The term “HAB” also applies to non-toxic blooms of macroalgae (seaweeds), which can cause major ecological impacts such as the displacement of indigenous species, habitat alteration and oxygen depletion in bottom waters.Globally, the nature of the HAB problem has changed considerably over the last several decades. The number of toxic blooms, the resulting economic losses, the types of resources affected, and the number of toxins and toxic species have all increased dramatically. Some of this expansion has been attributed to storms, currents and other natural phenomena, but human activities are also frequently implicated. Humans have contributed by transporting toxic species in ballast water, and by adding massive and increasing quantities of industrial, agricultural and sewage effluents to coastal waters. In many urbanized coastal regions, these inputs have altered the size and composition of the nutrient pool which has, in turn, created a more favorable nutrient environment for certain HAB species. The steady expansion in the use of fertilizers for agricultural production represents a large and worrisome source of nutrients in coastal waters that promote some HABs.The diversity in HAB species and their impacts presents a significant challenge to those responsible for the management of coastal resources. Furthermore, HABs are complex oceanographic phenomena that require multidisciplinary study ranging from molecular and cell biology to large-scale field surveys, numerical modelling, and remote sensing from space. Our understanding of these phenomena is increasing dramatically, and with this understanding comes technologies and management tools that can reduce HAB incidence and impact. Here I summarize the global HAB problem, its trends and causes, and new technologies and approaches to monitoring, control and management, highlighting molecular probes for cell detection, rapid and sensitive toxin assays, remote sensing detection and tracking of blooms, bloom control and mitigation strategies, and the use of large-scale physical/biological models to analyze past blooms and forecast future ones.     

有害赤潮对浮游动物摄食的影响

藻华是指水体中藻类大量繁殖的一种现象,有时这种浮游植物的爆发性增殖会造成海洋生物死亡,生态环境恶化,通过藻毒素污染海洋产品,从而对人类健康甚至生命构成威胁,国际科技界目前称这类藻华（赤潮）为有害藻华（harmful algal blooms HABs）,也称有害赤潮。近年来,有害赤潮爆发的频率和规模都在逐渐增加,造成的危害越来越大,赤潮现象有在全球蔓延的趋势。赤潮对海洋生态系统造成的影响越来越受到关注,世界各国也相应展开了研究。     

Increased diversity and environmental threat of harmful algal blooms in the Southern Yellow Sea,China

Harmful algal blooms(HABs) in the Southern Yellow Sea(SYS) have shown a trend of increasing diversity and detrimental ef fects. While the Bohai Sea, East China Sea, and South China Sea have experienced a high incidence of HABs since the 1980s, the Yellow Sea provides a relatively healthy ecological environment in which fewer HABs have been documented before the 21s t century. Yet largescale blooms of the green macroalga Ulva prolifera(so-called “green tides”) have occurred annually since 2007 in...     

Ecological features of harmful algal blooms in coastal upwelling ecosystems

Upwelling regions are the most complex habitats in which dinoflagellates produce red tides, but the flora is not unique. Many species also bloom in nutrient-enriched, non-upwelling systems, share the collective dinoflagellate trait of low-nutrient affinity, and can achieve relatively fast growth rates. Blooms occur over the range of nutrient – mixing – advection combinations found in upwelling habitats, rather than being restricted to the high-nutrient high-irradiance low-turbulence conditions posited by Margalef's classical Mandala and its Bowman et al. and Pingree versions. The bloom species are primarily ruderal strategists (R-species), which typify "mixing – drift" life-forms adapted to the velocities associated with frontal zones, entrainment within coastal currents, and vertical mixing during upwelling relaxations. Collectively, dinoflagellates appear capable of surviving fairly high turbulence spectra formed at representative Kolmogorov length scale – wind speed conditions. This biophysical protection might be the result of cell size-facilitated entrainment within the micro-eddies formed during turbulent energy dissipation. The swimming speeds of 71 clones of dinoflagellates are compared and related to reported rates of vertical motion in coastal upwelling systems. There are slow and fast swimmers; many exhibit motility rates that can exceed representative in situ vertical and horizontal water mass movements. At least four dinoflagellates from upwelling systems form chains leading to increased swimming speeds, and may be an adaptation for growth in coastal upwelling habitats. Red tides are frequent and fundamental features of upwelling systems, particularly during intermittent upwelling relaxations, rather than dichotomous (sometimes catastrophic) interruptions of the diatom blooms characteristically induced by upwelling. Successional sequences and the "red tide" zone may differ between upwelling and non-upwelling systems. In the latter, red tides diverge from the main sequence and are appropriately positioned in the Mandala's ecological space of high nutrients and low turbulence. An amended Mandala based on Pingree's S-kh model and the Smayda and Reynolds life-form model is presented to accommodate the range of red tide development and their successional routing found in coastal upwelling systems. Ecophysiological data support the Pitcher and Boyd seeding mechanism model, which can lead to red tides in upwelling systems. Nutrients, phyto-stimulation and grazing pressure as triggering factors in upwelling-system red tides are considered. Some red tides may be stimulated by nutrients and growth promoting factors excreted by migrating shoals and "boils" of c1upeoid stocks, with selective zooplankton grazing contributory. Substantial collapses in grazing pressure may be essential in anoxic red tide events. The mass mortalities that accompany anoxia, common to the Benguela and Peru upwelling systems, may be a trophic control mechanism to maintain biogeochemical balance and regional homeostasis, which are vital to upwelling ecosystem dynamics. Some traditional concepts of phytoplankton ecology may not completely apply to dinoflagellate bloom events in coastal upwelling systems.     

浙江近岸海域赤潮发生的趋势与特点

根据历史记载和近年调查实测资料,概述了浙江近岸海域赤潮的历史与现状,分析了赤潮的特点与发生趋势,研究结果表明:浙江近岸海域各种不同类型藻类引发的赤潮频率均呈上升趋势,其中甲藻赤潮的发生次数上升尤为迅速,赤潮出现的规模越来越大,有毒赤潮记录次数也显著增加。探讨了赤潮发生的一些原因,指出了建立科学评估有害赤潮影响体系和预警系统的意义并提出了一些对策措施与建议。     

Monitoring of harmful algal blooms along the Norwegian Coast using bio-optical methods

A Norwegian monitoring system for harmful algal blooms, consisting of an Observer Network, the State Food Hygiene Control Agency, the Oceanographic Company of Norway, the Institute of Marine Research and the Directorate for Fisheries, is reviewed. Potentially harmful algae on the Norwegian coast are found primarily in four classes of phytoplankton, dinoflagellates, prymnesiophytes, diatoms and raphidophytes. The system consists of buoys designed for real-time, in situ monitoring and forecasting, and is used principally to provide an early warning to the aquaculture industry. The system allows detection of potentially toxic species, through a combination of physical, chemical, biological and bio-optical data. New datasets using bio-optical techniques on glass-fibre filters are also described, providing information on the composition and health of phytoplankton populations.     

象山港水体的磷酸盐及其对赤潮的潜在影响

根据2012年5月和2017年10月在象山港海域的调查结果,研究了象山港磷酸盐的来源、分布和消耗。磷酸盐浓度在两次的调查中均呈现由港顶向湾口逐渐降低的趋势,说明港顶区有较高浓度磷酸盐的输入,可能主要来自生活污水和海水养殖。磷酸盐、硝酸盐和盐度在象山港外湾的分布特征说明长江和钱塘江是外湾水体磷酸盐的一个重要来源;此外,磷酸盐、硝酸盐和盐度在牛鼻山水道和佛渡水道的差异分布说明,陆地径流可能经由佛渡水道向象山港贡献了一部分的磷酸盐。内湾水体的磷酸盐分布受控于内外湾水体混合和浮游植物的同化作用,磷酸盐在水体停留时间较长的内湾发生了明显的消耗,相比于保守混合模型的估算值,磷酸盐在象山港内湾中的净消耗比例在2012年5月和2017年10月分别约为20%和9%。2017年秋季观测到的磷酸盐浓度[（1.88±0.31）μmol/L]与2002-2005年在同季节观测到的结果相比增加超过了50%,N/P值下降明显,这一变化可能会导致象山港内引发藻华的藻类种群发生变化,同时也可能会引发更为严重的赤潮。     

闽三角海域赤潮发生的时空特征

赤潮是海洋生态系统异常的重要指标。本文收集1986～2017年闽三角海域的赤潮事件,基于GIS技术分析赤潮的时空分布特征。结果表明:32年来,闽三角海域赤潮发生98次,赤潮高发区在厦门西海域、同安湾和泉州湄洲湾海域;闽三角海域赤潮年际发生次数呈波动变化,一年中赤潮高发期集中在5～8月;中肋骨条藻、角毛藻及旋链角毛藻是闽三角海域的赤潮优势种,具有一定的区域分布特征;有毒赤潮事件发生频率有增加趋势。本文系统性地总结了闽三角海域赤潮时空特征,为该区域的赤潮风险分析和预防提供依据。     

Physical processes contributing to harmful algal blooms in Saldanha Bay,South Africa

The study synthesises current understanding of the predominant physical processes responsible for the seasonality of harmful algal blooms, notably Alexandrium catenella and Dinophysis spp., in the nearshore environment of Saldanha Bay on the west coast of South Africa. Saldanha Bay is one of the few naturally sheltered areas on the South African coastline suitable for in situ shellfish farming and is the major site for the production of black mussel Mytilus galloprovincialis in South Africa. Mussel farming started there in 1985 and the present level of production is some 2 700 tons per annum. Since 1994, disruption of harvesting as a result of the presence of harmful algal species has been a regular late-summer phenomenon. Toxic blooms that are ultimately advected into the bay develop on the continental shelf to the north between 32°S and St Helena Bay, a region characterized by favourable conditions for dinoflagellate growth and circulation patterns that facilitate build-up of intense blooms during late summer. Offshore dinoflagellate populations are advected shorewards and polewards in response to relaxation of upwelling at the Namaqua cell to the north. Dinoflagellate blooms are advected south from the southern Namaqua shelf during upwelling relaxation. Under such conditions, the gyre south of Elands Bay moves offshore and a barotropic flow past Cape Columbine is established. Evidence suggests that the near surface component of the flow occurs as a sudden "flood" event. These dinoflagellate-containing shelf waters are in turn advected into Saldanha Bay when upwelling relaxes, when the density gradient between the bay and the shelf drives surface inflow and bottom water outflow. These flows are reversed with the resumption of upwelling over the shelf, resulting in intrusion and entrainment of bottom water and surface outflow. Entrainment dictates that the bay acts as a net importer of bottom water and net exporter of surface waters over a synoptic cycle. This system of exchange between Saldanha Bay and the shelf curtails the duration and severity of toxic episodes in the bay relative to the shelf.     

南海赤潮的时空分布特征及其与南海环境关系的综合分析

利用历史数据和卫星观测,系统分析20多年来(1980年-2003年)在南海海域发生的赤潮事件与环境特征,详细比较赤潮在南海四大区域(北部、东部、南部和西部海区)的区域分布、多发藻种和季节变化等特征。研究结果表明南海海域赤潮发生具有以下特点:(1)区域性:赤潮多发区域集中在珠江口附近(中国),马尼拉湾和马辛洛克湾(菲律宾)和沙巴州的西海岸(马来西亚);(2)季节性:在北部海区,赤潮的发生时间主要集中在3-5月份,东部海区在5-7月份,西部海区在7月份,而南部海区在全年都会发生;(3)藻种差异性:主要赤潮藻种在北部海区是夜光藻(Noctiluca scintillans),东部海区和南部海区是扁甲藻(Pyrodinium bahamense);(4)年际变化:不同海域赤潮的多发年份不同,对于整个南海而言,1991-1998年是赤潮的多发年。赤潮发生受到环境条件的影响,如季风、河流的排放、上升流、以及沿岸海域水体富营养化。     

A numerical model study on multi-species harmful algal blooms coupled with background ecological fields

Based on systematized physical, chemical, and biological modules, a multi-species harmful algal bloom （HAB） model coupled with background ecological fields was established. This model schematically embod-ied that HAB causative algal species and the background ecological system, quantified as total biomass, were significantly different in terms of the chemical and biological processes during a HAB while the inter-action between the two was present. The model also included a competition and interaction mechanism between the HAB algal species or populations. The Droop equation was optimized by considering tempera-ture, salinity, and suspended material impact factors in the parameterization of algal growth rate with the nutrient threshold. Two HAB processes in the springs of 2004 and 2005 were simulated using this model. Both simulation results showed consistent trends with corresponding HAB processes observed in the East China Sea, which indicated the rationality of the model. This study made certain progress in modeling HABs, which has great application potential for HAB diagnosis, prediction, and prevention.     

Trophic cascades and future harmful algal blooms within ice-free Arctic Seas north of Bering Strait: A simulation analysis

Within larger ice-free regions of the western Arctic Seas, subject to ongoing trophic cascades induced by past overfishing, as well as to possible future eutrophication of the drainage basins of the Yukon and Mackenzie Rivers, prior very toxic harmful algal blooms (HABs) – first associated with ∼100 human deaths near Sitka, Alaska in 1799 – may soon expand. Blooms of calcareous coccolithophores in the Bering Sea during 1997–1998 were non-toxic harbingers of the subsequent increments of other non-siliceous phytoplankton. But, now saxitoxic dinoflagellates, e.g. Alexandrium tamarense, were instead found by us within the adjacent downstream Chukchi Sea during SBI cruises of 2002 and 2003. A previous complex, coupled biophysical model had been validated earlier by ship-board observations from the Chukchi/Beaufort Seas during the summer of 2002. With inclusion of phosphorus as another chemical state variable to modulate additional competition by recently observed nitrogen-fixers, we now explore here the possible consequences of altered composition of dominant phytoplankton functional groups [diatoms, microflagellates, prymnesiophyte Phaeocystis colonies, coccolithophores, diazotrophs, and dinoflagellates] in relation to increases of the toxic A. tamarense, responding to relaxation of grazing pressure by herbivores north of Bering Strait as part of a continuing trophic cascade. Model formulation was guided by validation observations obtained during 2002–2004 from: cruises of the SBI, CHINARE, and CASES programs; moored arrays in Bering Strait; other RUSALCA cruises around Wrangel Island; and SBI helicopter surveys of the shelf-break regions of the Arctic basin. Our year-long model scenarios during 2002–2003 indicate that post bloom silica-limitation of diatoms, after smaller simulated spring grazing losses, led to subsequent competitive advantages in summer for the coccolithophores, dinoflagellates, and diazotrophs. Immediate top-down control is exerted by imposed grazing pressures of the model’s herbivores and bottom-up control is also effected by light-, nitrate-, ammonium-, silicate-, and phosphate-modulated competition among the six functional groups of the simulated phytoplankton community. Similar to the history of the southern North Sea adjacent to the Rhine River, possible farming of northwestern Alaska and Canada, in conjunction with other human activities of ice retreat and overfishing, may lead to future exacerbations of poisonous phytoplankton. These potential killers include both toxic dinoflagellate and diazotroph HABs, deadly to terrestrial and marine mammals, as well as those of prymnesiophytes, some of which have already foamed beaches, while others have killed fishes of European waters.     

Numerical investigation of the effects of upwelling on harmful algal blooms off the west Florida coast

The initiation of the toxic harmful algal bloom (HAB), Karenia brevis, along the west Florida coast has been associated with upwelling events. Upwelling processes may be responsible for the transport of nutrients or algae from deep offshore locations across the Florida shelf to the coast. The influence of coastal wind-driven upwelling on the onset and occurrences of K. brevis in this region was numerically investigated using Rutgers University's Regional Ocean Modeling System. Computations were carried out in an idealized model domain, a two-dimensional slice in the cross-shore and vertical directions. The surface forcing data used was from several offshore meteorological buoys. The motion of the algae was simulated using Lagrangian particles and a passive tracer. The numerical simulations of three K. brevis events in 2000–2002 showed that the particles respond (with upwelling/downwelling) to the along-shore wind stresses as expected and some upwelling was present during the events. Comparison of the passive tracer fields with measured fluorescence data exposed the model's sensitivity to the particular surface forcing data employed and the relatively more significant role played by surface forcing over initial conditions. The present model set-up constitutes a useful predictive tool for conditions conducive to the onset of HABs. It is planned to be used in a real-time mode to aid the NOAA HAB monitoring and forecasting system.     

赤潮的高光谱监测及预警方法

有害赤潮发生频度有逐年增加的趋势, 建设赤潮的早期监测和预警系统是一个难点。文章研究表明, 珠江口赤潮前后, 叶绿素a浓度和细胞密度变化范围分别为0.9—31.1mg•m-3 和1.28×104 —1.76×106cell•L-1, 优势藻为聚生角毛藻; 高光谱辐射计获取的光漫射衰减系数、遥感反射率和荧光强度可反演叶绿素a浓度和细胞密度, 其中反演叶绿素a浓度的平均相对均方根差(RRMS)为30.6%、33.8%和77.4%, 而反演细胞密度的平均RRMS为83.6%、83.9%和136.4%。因高光谱辐射计可以获取每小时或者更短时间尺度的数据, 因此, 装备了高光谱辐射计的光学浮标可用于赤潮监测与预警系统。     

2005年春夏季东海赤潮过程中营养盐作用初探

自20世纪90年代以来,我国进入赤潮高发期,近岸海域赤潮的发生频率、波及范围和危害程度呈逐年上升趋势.受长江径流等陆源输入影响,东海海域形成了其特有的富营养环境,目前已经成为我国赤潮发生最严重的区域之一.该海域赤潮暴发呈现出发生时间不断提前、持续时间不断加长、发生面积不断扩大、有毒有害藻类逐渐增加的特点[1-2].据统计,自2000年以来,我国沿岸海域共发生赤潮约480次,累计面积超过104000km2.其中,仅在东海海域发生赤潮的次数就占全国记录总数的60%,发生面积更是达到了70%以上[3],其发生频率和规模均远远高于我国其他3个海区.     

Short-term and diurnal temperature changes alter the response of harmful algal blooms of Pseudo-nitzschia pungens to solar ultraviolet radiation

In this study, we explored the interactive effects of temperature and solar ultraviolet radiation (UVR) on the growth, pigment contents, photochemical efficiency and non-photochemical quenching (NPQ) of Pseudo-nitzschia pungens, which forms harmful algal blooms globally. Cells were first pre-acclimated to temperatures of 20°C, 25°C or 28°C for 10 days. They were then exposed to three different types of solar radiation for 60?min. We then measured the photochemical efficiency of the cells during the 60-min exposure and a 360-min recovery period under dim light (20?μmol?photons?m^?2?s^?1). In addition, we analysed the diural change in photochemical efficiency and NPQ over a 10-h period. We found that P. pungens that were exposed to higher temperatures in the short term (i.e. 20–25°C, 20–28°C and 25–28°C) or long term (i.e. 25–25°C and 28–28°C) exhibited lower levels of photoinhibition and faster recovery rates than samples exposed to a lower temperature (i.e. 20–20°C). This indicated that the deleterious effects of UVR can be minimised by both chronic and acute exposure to higher temperatures within the tolerated temperature range for P. pungens.     

春季东海有害藻华高发区仔稚鱼时空分布

根据2011年春季(3月29日—5月27日)海洋综合调查,对东海有害藻华高发水域(28°15′36″~30°30′16″N、121°52′36″~123°15′6″E)的仔稚鱼种类组成、时空分布以及主要优势种与环境因子的相关性进行了探讨。调查共采集到仔稚鱼27种,前几位优势种分别为日本鳀Engraulis japonicus、鰕虎科sp.(Gobiidae sp.)、六丝钝尾鰕虎Amblychaeturichthys hexanema、小黄鱼Larimichthys polyactis,该4种占据了总丰度的91.23%。优势种丰度和发育阶段组成的时空变化显示,日本鳀在调查期间有两次产卵小高峰,其仔鱼孵化后向岸漂流;小黄鱼和六丝钝尾鰕虎在此期间只有一次产卵高峰,其仔鱼在舟山群岛邻近水域进行保育。可以看出,调查水域及邻近水域是部分鱼类产卵,仔稚鱼保育、向岸漂流的场所或途径,是鱼类早期生活史中重要生境,一旦暴发有害藻华,补充群体将受到不良影响。多元分析结果显示,所选环境因子对几位优势种仔稚鱼的分布影响不大,推测产卵场位置和保育方式可能是更重要的原因。