棕囊藻的生物学概述 I.形态分类和生理生态学

海洋浮游植物棕囊藻Phaeocystisspp.在自然界常以游离单细胞和由粘胶质包裹的细胞集群体两种细胞形式轮换出现,它在世界各营养丰富的海域中无处不在,并可形成高细胞密度、暴发性繁殖及形成赤潮。rRNA基因研究数据说明,目前全世界海洋分布的Phaeocystisspp.有3个分开种,即Phaeocystispouchetii、Phaeocystisglobosa和Phaeocystisantarctica。棕囊藻奇特的生理学和生态学特征是独一无二的,它复杂的多形态变化生命循环导致海域植物性和海底食物网(如微量元素的生物地球化学)的结构与功能均发生了极大的变化。棕囊藻的生命循环体现了其主要的生态学特征,不同棕囊藻形态的生物化学和营养代谢与细菌,微型、中型和后生浮游动物的作用以及海洋环境中的物理结构均有关。在自然界,棕囊藻的生命循环中最常见和最重要的是它的集群体形态,集群体形态是一个具有整体生物学功能特征的生物实体。棕囊藻在生态系中起到重要作用的关键在于集群体形态在合成糖粘胶质的能力较强。     

棕囊藻的生物学概述—I．形态分类和生理生态学

海洋浮游植物棕囊藻Phaeocystis spp.在自然界常以游离单细胞和由粘胶质包裹的细胞集群体两种细胞形式轮换出现,它在世界各营养丰富的海域中无处不在,并可形成高细胞密度、暴发性繁殖及形成赤潮.rRNA基因研究数据说明,目前全世界海洋分布的Phaeocystis spp.有3个分开种,即Phaeocystis pouchetii、Phaeocystis globosa和Phaeocystis antarctica.棕囊藻奇特的生理学和生态学特征是独一无二的,它复杂的多形态变化生命循环导致海域植物性和海底食物网(如微量元素的生物地球化学)的结构与功能均发生了极大的变化.棕囊藻的生命循环体现了其主要的生态学特征,不同棕囊藻形态的生物化学和营养代谢与细菌,微型、中型和后生浮游动物的作用以及海洋环境中的物理结构均有关.在自然界,棕囊藻的生命循环中最常见和最重要的是它的集群体形态,集群体形态是一个具有整体生物学功能特征的生物实体.棕囊藻在生态系中起到重要作用的关键在于集群体形态在合成糖粘胶质的能力较强.     

不同氮源对球形棕囊藻生长和囊体形成的影响

棕囊藻引发的有害藻华已经成为一种全球性的自然灾害,给海洋环境以及渔业经济带来了严重的损失。棕囊藻存在两种生活史形态:游离的单细胞形态和囊体形态,但是不同形态之间的转换和囊体形成机制尚不清楚。营养盐是浮游植物生长和藻华发生的物质基础,棕囊藻藻华以往多发生在硝酸盐限制海区。但是近年来,尿素等有机氮在河口区域浓度不断提高,有机氮源的增加可能对棕囊藻生长和生活史转换产生显著的影响。通过添加硝酸盐、铵盐、尿素三种氮源,研究不同氮源对球形棕囊藻生长和囊体形成的影响。研究表明:球形棕囊藻可以在硝氮、尿素中迅速生长,形成囊体,但是氨氮对其却有明显的抑制作用;球形棕囊藻单细胞在硝氮中丰度比较高,最大可达到(444.21±64.97)×10~3个·mL~(-1);在尿素中更容易形成囊体,最多能达到(12.61±6.5)个·mL~(-1)。球形棕囊藻可以利用硝酸盐和有机氮源使其具有更强的竞争力,尿素在海洋水体中浓度的提高,可能是近年来球形棕囊藻藻华在我国近海水域频发的原因之一。     

小角毛藻对球形棕囊藻囊体形成的影响

球形棕囊藻具有异型的生活史,能够在单细胞和囊体两种不同形态之间转换。球形棕囊藻藻华现场发现,囊体有时附着在角毛藻上。为研究角毛藻对囊体形成的影响和机制,进行球形棕囊藻和小角毛藻的共培养实验,观察球形棕囊藻囊体的形成和生长状况,研究小角毛藻对球形棕囊藻囊体形成的影响。结果证明:混合培养下,小角毛藻和球形棕囊藻的囊体附着在一起,在较短的生存周期内,囊体的密度明显高于单独培养时的密度;而单独培养体系下,棕囊藻单细胞密度和囊体体积均高于混合培养,囊体的存在时间较长。小角毛藻的存在对囊体的形成具有促进效应,这对于球形棕囊藻囊体具有重要的生态意义。硅藻的存在更能体现球形棕囊藻竞争优势。     

不同摄食压力下球形棕囊藻的防御策略

球形棕囊藻是我国沿海常见有害藻华物种,生活史中包括囊体和单细胞两种形态,藻华爆发时的优势形态是囊体,游离单细胞很少被发现,但是球形棕囊藻囊体形成的机制目前还不清楚。本研究通过分析桡足类、纤毛虫和异养甲藻及其化学信号对囊体形成的影响,以探究摄食压力对球形棕囊藻生活史转变的影响。实验结果表明:3种摄食者的直接摄食均引发了球形棕囊藻囊体直径的扩大,摄食者的粒径越大,球形棕囊藻的囊体直径增大越显著,安氏伪镖水蚤的化学信号也能引发球形棕囊藻囊体显著增大,但是游仆虫和海洋尖尾藻的化学信号并未引发类似响应。囊体形成和直径的扩大保护了囊体内细胞免受摄食,有助于球形棕囊藻藻华的发生。     

中国沿海球形棕囊藻(Phaeocystis globosa)的分类、分布及其藻华

球形棕囊藻(Phaeocystis globosa Scherffel)是全球海洋广泛分布的有害藻华种类。1997年10月,中国东南沿海首次暴发了此种藻类的大规模藻华,其后陆续在福建、广东、广西、海南、河北及天津等省市沿海暴发多起同种藻华。中国近海的球形棕囊藻藻华呈现两大独有的特点,即藻类囊体较大(可达3厘米),以及藻华可毒害养殖业。历经20多年,球形棕囊藻在中国沿海已从一个"藻华新记录种"变成了"藻华常见种"。值得注意的是, 2014年以来,广西北部湾海域棕囊藻藻华肆虐,威胁核电冷源安全,受到了社会高度关注,也对球形棕囊藻藻华的研究提出了新的挑战与要求。针对这一生态灾害的最新发展趋势,本文总结了20年来中国球形棕囊藻及其藻华灾害的发生与发展状况,分别就棕囊藻的分类、生活史特征、营养特性、藻华形成的环境驱动因素、生态毒理等诸多方面开展简要综述,冀望为棕囊藻藻华的研究及防治提供基础资料及思路。     

中国新记录定鞭藻物种冠状棕囊藻的遗传进化分析

棕囊藻属(Phaeocystis)分类学上位于定鞭藻门(Haptophyta)定鞭藻纲(Haptophyceae)棕囊藻目(Phaeocystales)棕囊藻科(Phaeocystaceae)。迄今在我国海域仅分离鉴定到一种棕囊藻属物种,即自1997年以来在我国海域频繁形成有害藻华的球形棕囊藻(Phaeocystis globosa)。作者从中国南海分离到一个单细胞鞭毛类株系CNS01077,结合形态特征观察及基于18S rDNA序列的系统发育分析,将其鉴定为冠状棕囊藻(Phaeocystis rex)。这是该棕囊藻物种在我国海域的首次报道。该研究构建了该物种的首个叶绿体基因组序列和首个线粒体基因组序列。与球形棕囊藻和南极棕囊藻的细胞器基因组比较分析发现冠状棕囊藻的细胞器基因组发生了显著的结构重排和序列变异。该物种在我国海域的发现及细胞器基因组的构建,将为棕囊藻的生物多样性组成和地理分布研究提供支撑。     

北部湾棕囊藻藻华原因种分析

近年来, 棕囊藻已成为我国北部湾海域藻华发生的主要原因种, 严重影响北部湾海洋生态环境、水产增养殖业和防城港核电安全, 急切需要对棕囊藻藻华原因种作进一步分析确认。2017年2月和3月在钦州湾和涠洲岛藻华发生海域分离得到9株棕囊藻, 采用光学显微镜进行形态学初步鉴定, 并通过核糖体小亚基序列(SSU rDNA)进行系统进化分析。研究结果表明, 北部湾9株棕囊藻与球形棕囊藻形态特征基本符合, 系统发育树与不同海域来源的球形棕囊藻聚在同一大分支上, 与其他海域来源的球形棕囊藻遗传距离为0.0000~0.0071, 其中北部湾WZS3-1株与墨西哥湾株CCMP627、苏里南株CCMP628、厄瓜多尔株CCMP1528、南非株P162、中国海株RCC:K1398及汕头株Santou97的球形棕囊藻亲缘关系最近, 遗传距离为0.0000, 而北部湾WZS1、WZS2、WZS4、WZS5株与香港株SKLMP_T005和渤海株BOHAI1亲缘关系最近, 遗传距离为0.0000。球形棕囊藻的种内遗传距离(0.0000~0.0073)明显小于种间遗传距离(0.0084~0.0440), 因此可确定来自北部湾不同海域的9株棕囊藻均为球形棕囊藻。此外, 球形棕囊藻各藻株间的亲缘关系远近与其地理位置分布不能完全对应。本研究结果将为我国棕囊藻藻华藻种库构建、棕囊藻北部湾株生理生态研究和北部湾棕囊藻藻华发生机理研究等提供参考。     

紫外辐射对南极棕囊藻细胞DMSP合成和DMS释放率的影响

在不同紫外辐射波段下,南极棕囊藻(Phaeocystis antarctica)细胞的生长率、叶绿素a、细胞内DMSP含量和DMS释放量变化的测定结果表明;UV-B对南极棕囊藻细胞生长率和叶绿素a含量有抑制效应,UV-B还可加快DMSP分解成DMS和丙烯酸的分解速率,而UV-A对该藻细胞的DMSP合成有强烈的抑制效应。鉴于在每年春季极地海洋浮游植物繁殖期间,南极棕囊藻在南极海冰带海洋浮游植物种群结构中占有的优势地位,以及该藻是极地海洋浮游植物中DMS的主要释放者,推测南极“臭氧空洞”所增加的紫外辐射可能会对南极海域的DMS释放率产生一定的影响。     

四烷铵络合碘对球形棕囊藻去除作用的研究

研究了四烷铵络合碘对赤潮生物球形棕囊藻Phaeochystis globosa的灭杀和控制作用。结果表明,四烷铵络合碘对球形棕囊藻具有明显的抑制和杀灭作用。当藻细胞起始密度为1.5×106cells.ml-1时,低于15 mg.L-1的四烷铵络合碘溶液对球形棕囊藻生长有一定的抑制作用,高于25 mg.L-1的四烷铵络合碘溶液对球形棕囊藻有显著的杀灭作用。四烷铵络合碘具有高效、低毒、药效期长等优点,可能是一种较理想的除藻剂。     

Molecular identification of bloom-forming species Phaeocystis globosa (Pryninesiophyta) and its dispersal based on rDNA ITS sequence analysis

The colony-forming Phaeocystis species are causative agents of dense bloom occurrences in coastal waters worldwide. It is difficult to separate them because of the different morphologies associated with their colonial stages. In this study we applied molecular approaches to analyze the genetic variation of Phaeocystis globosa and Phaeocystis pouchetii from several geographic regions, and to assist in tracing the dispersal of bloom-forming Phaeocystis species in coastal waters of China. The sequences of the internal transcribed spacers (ITS1 and ITS2) of rDNA and the 5.8S ribosomal RNA gene of Phaeocystis strains were determined. Sequence comparison shows that P.globosa was the most divergent to P. pouchetii, exhibiting sequence divergence higher than 0.08. However, lower genetic divergences existed between strains of P.globosa. The sequence comparison of the Phaeocystis rDNA ITS clearly shows that the species isolated from the southeast coast of China is identified as P.globosa rather than P. cf. pouchetii or other species. Furthermore, the significance of rDNA variation in distinct global populations of P.globosa suggested it might have had sufficient time to accumulate detectable mutations at the rDNA locus, supporting the hypothesis of ancient dispersal of P.globosa to many areas, meaning that P.globosa blooms in the coastal waters of China are endemic rather than a newly introduced species or a foreign source. Finally, based on the high divergent region of rDNA ITS, a pair of species-specific primers for P.globosa were designed, they could be useful to detect the presence of this species in mixed plankton assemblages or flagellate stages that are recognized with diffculties by means of conventional microscopy.     

Protoraphis Simonsen,a newly recorded marine epizoic diatom genus for China

Epizoic diatoms on marine copepods are common in nature and may have a special ecological relationship with their hosts.However,this special ecological group is not well known,and it has only rarely been studied in the China seas.To address this knowledge gap,the species diversity and classification of epizoic diatoms on planktonic copepods were studied with samples collected from the East China Sea.In the present study,a marine araphid diatom genus Protoraphis and its type species,Pr.hustedtiana,were observed and identified by light and electron microscopy,thus representing the first record of this genus and its type species in China.This genus is characterized by a median sternum strongly bent to opposite sides and terminate in two transapical grooves at the valve ends.Protoraphis hustedtiana was found to be epizoic on the posterior body appendages and segments of the marine calanoid copepod Candacia bradyi.An internal view shows a complex,ear-shaped process that is close to the apical slit field.The ecological habitats and geographical distributions of Protoraphis were also discussed,and,together with complementary morphological studies,our results have increased the number of records for marine epizoic diatoms to three genera with three species in China,including Pseudohimantidium and Pseudofalcula.     

Morphological observation and rbcL gene sequences studies of two new species, Grateloupia dalianensis H.W.Wang et D.Zhao, sp. nov. and G. yinggehaiensis H.W.Wang et R.X.Luan, sp. nov. (Halymeniaceae, Rhodophyta) from China

A few species in the genus Grateloupia (Halymeniaceae,Rhodophyta) have been investigated in detail with respect to morphological observations and molecular analyses.In this study,the authors document the vegetative and reproductive structures of two new species of Grateloupia,G.dalianensis H.W.Wang et D.Zhao,sp.nov.and G.yinggehaiensis H.W.Wang et R.X.Luan,sp.nov.They both have the morphological character that carpogonial ampullae and auxiliary cell ampullae are the simple Grateloupia-type.The two species can be distinguished from other species of the genus by their distinctive morphological features respectively.Based on ribulose-1,5-bisphosphate carboxylase/oxygenase (rbcL) gene sequences,the phylogenetic tree obtained in the study indicated that they are both embedded within the Grateloupia clade.G.dalianensis clusters a subclade with G.asiatica,and G.yinggehaiensis forms a single monophyletic subclade with G.hawaiiana.     

棕囊藻赤潮原因种的分子鉴定和起源分析

测定了发生于我国东南海域赤潮原因种Phaeocystis sp.及相关种类P.globosa和P.pouchetii 18S rDNA序列,并以Phylip35分析软件构建序列距离矩阵和分子系统发育树图.结果表明,该赤潮原因种与球形棕囊藻P.globosa 18S rDNA序列完全相同,从分子水平鉴定了该原因种为P.globosa.分子系统发育树图表示的P.globosa和P.pouchetii的分化顺序表明,分布于我国东南海域的P.globosa可能是一种本地起源的暖水种.Phaeocystis globosa不同株系的形态差异,可能是适应不同生活环境的结果.根据分子生物学的数据对有害赤潮藻类进行系统发育分析,并设计用于有害赤潮藻快速鉴定的特异性分子探针,对赤潮的监控和防治具有重要意义.     

有害赤潮藻赤潮异弯藻Heterosigma akashiwo研究进展

浮游植物不仅在食物链中充当贡献者,也在生物地球化学循环过程中扮演着重要角色,然而一些种类在某些条件下暴发性增殖或聚集,形成有害藻华,对人类健康和生态系统构成威胁。赤潮异弯藻是形成有害藻华的典型物种,其藻华发生时会造成养殖鱼类大面积死亡,给水产养殖业造成巨大损失。近年来,随着全球气候变暖、水体富营养化程度日趋严重,有害藻华发生的规模和频次总体呈增加趋势,且相关研究表明气候变暖和二氧化碳水平升高均能刺激赤潮异弯藻的生长。本文主要从赤潮异弯藻物种鉴定、基础生物学研究、生态学研究等几个方面综述了该物种的研究进展,并对其未来研究方向进行了相关展望,以期为赤潮异弯藻的藻华形成机制研究、藻华防治等工作提供参考。     

Grazing experiments and model simulations of the role of zooplankton in Phaeocystis food webs

A combined empirical and modelling study was conducted to further examine the potential importance of grazing by zooplankton in pelagic food webs in which Phaeocystis is a significant or dominant component. Laboratory experiments were designed to measure ingestion of Phaeocystis and other potential prey items which co-occur with Phaeocystis. Grazers included copepods and ciliates, and prey included Phaeocystis colonies and solitary cells, diatoms, ciliates, bacteria, and detritus. These data were expressed in the model currency of nitrogen units, and fit to hyperbolic tangent equations which included minimum prey thresholds. These equations and literature data were used to constrain a food web model whose purpose was to investigate trophic interactions rather than to mimic actual events. Nevertheless, the model output was similar to the general pattern and magnitude of development of Phaeocystis–diatom communities in some environments where they occur, e.g. north Norwegian waters. The model included three forms of nitrogen, three phytoplankton groups, bacteria, two zooplankton groups, and detritus, with detailed flows between compartments. An important component of the model was inclusion of variable prey preferences for zooplankton. The experiments and model simulations suggest several salient conclusions. Phaeocystis globosa colonies were eaten by a medium-sized copepod species, but ingestion appeared to be strongly dependent upon a proper size match between grazer and prey. If not, colonies were eaten little if at all. Phaeocystis solitary cells were ingested rapidly by ciliate microzooplankton, in agreement with prior literature observations. In contrast, detritus was eaten comparatively slowly by both ciliates and copepods. Both types of zooplankton exhibited apparent minimum prey thresholds below which grazing did not occur or was inconsequential. Model simulations implied that transitions between life cycle stages of Phaeocystis may potentially be important to phytoplankton–zooplankton interactions, and that relative rates of ingestion of Phaeocystis by various zooplankton may have significant impacts upon material fluxes through and out of Phaeocystis–diatom ecosystems. Indirect effects of trophic interactions appear to be equally significant as direct effects.     

中国海洋仙菜科(Ceramiaceae)新记录属——盖氏藻属Gayliella(红藻门)的分类研究

利用生物压片技术,通过形态学和解剖学方法,对采自我国沿岸海洋的仙菜科红藻(Ceramiaceae,Rhodophyta)进行了分类学研究,发现了我国仙菜科新记录属——盖氏藻属(新拟名)Gayliella T.O.Cho,L.McIvor et S.M.Boo。该属目前在我国共有5个种,即短毛盖氏藻(新拟名)Gayliella pmbriatum (Setchell et N. L. Gardner). T. O. Cho et S. M. Boo、优美盖氏藻(新拟名) Gayliella βaccidum (Kützing) T. O. Cho et L. McIvor、泰式盖氏藻(新拟名) Gayliella taylorii (E. Y. Dawson) T. O.Cho et S. M. Boo、马沙盖氏藻(新拟名) Gayliella mazoyerae T. O. Cho, Fredericq et Hommersand和横轴盖氏藻(新拟名) Gayliella transversalis (Collins et Hervey) T. O. Cho et Fredericq,...     

Production of methyl bromide and methyl chloride in laboratory cultures of marine phytoplankton II

Methyl halides (monohalomethanes), especially methyl bromide, are known to contribute significantly to ozone destruction in the stratosphere. Budgets of natural and anthropogenic methyl bromide suggest that marine organisms may be the source of a significant proportion of the total global production. Since phytoplankton are abundant in surface waters, they are obvious candidates. Cultures of nine phytoplankton species were grown in CO₂-enriched, nitrate-limited medium in sealed glass vessels. Species tested include Chaetoceros calcitrans, Isochrysis sp., Porphyridium sp., Synechococcus sp., Phaeodactylum tricornutum, Tetraselmis sp., Prorocentrum sp., Emiliania huxleyi and Phaeocystis sp. Methyl bromide (CH₃Br) and methyl chloride (CH₃Cl) concentrations were determined by bubbling the cultures with high-purity air, cryotrapping the effluent and analyzing it on a gas chromatograph with an electron capture detector. The Phaeocystis sp. samples were monitored with GCMS. Cell population, bacterial population, Chl a, pH, and nitrate concentration were monitored for periods of at least two weeks. CH₃Cl was produced by all cultures. CH₃Br was absent in Tetraselmis sp. and Isochrysis sp. cultures, but present in all the others. Methyl iodide (CH₃I) was present in most cultures but could not be quantified due to analytical limitations. CH₃I and CH₃Br production was fastest in stationary phase and continued long after cell division had ceased. Axeic cultures of Phaeocystis sp. achieved almost identical production rates of CH₃Cl and CH₃Br as xenic cultures of the same species. Species from tropical waters had faster CH₃Cl and CH₃Br production rates than temperate species. Scaling the observed production rates using global standing stock estimates for Chl a and particulate nitrogen indicate that phytoplankton can account for only a fraction of the CH₃Cl and CH₃Br believed to be produced in the ocean. Calculations based on the estimated global biomass of Phaeocystis sp. and E. huxleyi blooms show that their contribution is insignificantly small.