Relationship between bacteria and phytoplankton during the giant jellyfish Nemopilema nomurai bloom in an oligotrophic temperate marine ecosystem

Bacterial abundance, phytoplankton community structure and environmental parameters were investigated to study the relationships between bacteria and phytoplankton during giant jellyfish Nemopilema nomurai blooms in the central Yellow Sea during 2013. N. nomurai appeared in June, increased in August, reached a peak and began to degrade in September 2013. Results showed that phosphate was possible a key nutrient for both phytoplankton and bacteria in June, but it changed to nitrate in August and September. Phytoplankton composition significantly changed that pico-phytoplankton relative biomass significantly increased, whereas other size phytoplankton significantly decreased during jellyfish bloom. In June, a significantly positive correlation was observed between chlorophyll a concentration and bacterial abundance（r=0.67, P0.001, n=34）.During jellyfish outbreak in August, there was no significant correlation between phytoplankton and bacteria（r=0.11, P0.05, n=25）, but the relationship（r=0.71, P0.001, n=31） was rebuilt with jellyfish degradation in September. In August, small size phytoplankton occupied the mixed layer in offshore stations, while bacteria almost distributed evenly in vertical. Chlorophyll a concentration significantly increased from（0.42±0.056） μg/L in June to（0.74±0.174） μg/L in August, while bacterial abundance just slightly increased. Additionally, the negative net community production indicated that community respiration was not entirely determined by the local primary productivity in August. These results indicated that jellyfish blooms potentially affect coupling of phytoplankton and bacteria in marine ecosystems.

Asexual Reproduction of Aurelia coerulea Polyps in situ on Settling Plates in Jiaozhou Bay,China

Jellyfish blooms are increasingly prevalent coastal phenomena,with Aurelia coerulea being a significant contributor.The complex life history of jellyfish,featuring a sessile polyp stage,is a key factor in bloom formation,yet it is challenging to study in the field.This study aimed to describe the asexual reproduction of A.coerulea polyps in Jiaozhou Bay,China,and to assess their ability to form sedentary colonies and their relationship with environmental factors.Using settling plates,we observed polyps in situ and documented three asexual reproduction modes:budding,strobilation,and podocyst formation.Environmental parameters including temperature,salinity,dissolved oxygen(DO),chlorophyll a(Chl a),and the biomass of phytoplankton and zooplankton were monitored.Budding was the predominant reproductive method,featuring stolon budding in autumn and direct budding during the winter and spring seasons.Strobilation took place in the colder months,releasing ephyrae between April and May.Polyp abundance was positively correlated with seawater temperature and negatively correlated with DO concentration.Polyps at 1.0 m depth showed higher survivorship and proliferation over 8 months,whereas those at 2.0 m depth experienced a steady decline and died within 4 months.The study enhances our understanding of the life cycle and asexual reproduction strategies of A.coerulea polyps in the field,with implications for predicting and managing jellyfish blooms.The findings underscore the importance of temperature and DO in polyp survival and suggest that initial polyp population proliferation is crucial for long-term survival.

Intraguild predation by polyps of three scyphozoan jellyfish:Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum

Jellyfish blooms occur worldwide and have resulted in serious problems in tourism,fisheries,coastal industries,and the marine ecosystem.The life cycle of scyphozoan jellyfish consists of a pelagic medusa stage and a benthic polyp stage.Success of asexual reproduction of the polyps determines directly the number of medusae;thus,the polyp stage is the key to understanding the population dynamics of medusae.Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum are three scyphozoan jellyfish commonly inhabit in Chinese coastal waters.Polyps of A.coerulea are easily visible,while those of N.nomurai and R.esculentum remain yet to be found in the wild.However,distribution of the medusa indicates that the polyps of all three species may occur together.To evaluate the distribution pattern of polyps of the three species and explore intraguild predation by the polyps,we conducted a laboratory experiment that considered the attachment sequence and size relationship of calyx diameter of the polyps.We found that the polyps of A.coerulea preyed on polyps of the other two species in all treatments,except when polyps of R.esculentum were bigger than those of A.coerulea.The polyps of R.esculentum preyed on the polyps of N.nomurai only when polyps of R.esculentum attached first and were bigger than those of N.nomurai.Colonies of N.nomurai polyps were rarely found in the places inhabited by A.coerulea polyps.In addition,A.coerulea polyps are known to inhabit at depths of less than 20 m in coastal sea,thus,we speculate that N.nomurai polyp colonies might occur at depths of more than 20 m.Therefore,our finding that polyps of A.coerulea aggressively preyed on polyps of other species may help understand other such systems of jellyfish bloom in the world.

黄、东海水母暴发机理及其生态环境效应研究进展

在全球变化和人类活动影响下海洋生态系统的结构与功能发生了很大的变化,海洋赤潮、绿潮、白潮(水母暴发)等生态灾害在多重压力下不断出现。对于以水母为代表的胶质类生物的数量增多,国际上有很多的解释和争论:水母的暴发是一种趋势性的还是周期性的现象,关键问题是对导致水母暴发原因的认识。针对这些问题,国家重点基础研究发展计划项目"中国近海水母暴发的关键过程、机理及生态环境效应"开展了综合性的研究,对前期的研究进展和结果进行系统整理发表在本专辑中,涉及水母生活史策略、环境因子对水母不同生活史阶段的生长和发育的影响、水母的暴发对环境的影响、以及对中国近海水母暴发的新认知等。

水母灾害的形成机理、监测预测及防控技术研究进展

受人类活动和全球变化双重压力的影响,近年来我国近海水母呈现增多的趋势,局部海域水母灾害连年发生,对近海核电站等重大设施安全运行、海洋经济可持续发展和海洋生态环境安全造成严重威胁。我们不仅亟需在科学上阐明我国近海水母灾害发生的重要过程和机制,而且在技术上亟需研发水母灾害监测预警系统和防灾减灾技术体系,以提升我国海洋水母灾害应对能力和海洋环境安全保障能力。本文在查阅文献资料、国际交流与合作、课题组的研究成果的基础上,综述国内外水母灾害的形成机理、水母灾害监测及防控技术的研究进展,分析各种水母监测技术,防控技术的局限性以及在特定海域成功应用的原因,展望今后我国水母灾害的监测技术及防控策略。

环境因子对海月水母螅状体存活与繁殖的影响

以我国黄渤海主要致灾大型水母海月水母的螅状体为实验材料,采用实验生态学方法研究了温度、盐度、光照强度、饵料量对其存活与繁殖的影响。结果表明,在0—25℃范围内,螅状体40 d的成活率均为100%,螅状体相对繁殖速度随温度的升高而增加;盐度在15—40范围内,螅状体40 d的成活率均为100%,盐度22.5组的螅状体相对繁殖速度最高,但盐度对螅状体种群数量影响不显著;螅状体相对繁殖速度随光照强度的增加而下降,黑暗条件有利于海月水母螅状体个体数量增加;螅状体相对繁殖速度随饵料量的增加而增加。研究结果说明海月水母螅状体对环境具有较强的适应性,水温升高、饵料量增加可导致海月水母螅状体种群数量快速增加。

大型水母爆发对东海生态系统中上层能量平衡的影响

通过建立东海生态系统ECOPATH模型,并将大型水母作为一个独立的功能组,从能量平衡的角度探讨近年来东海大型水母爆发对生态系统的影响,并在此基础上提出抑制大型水母爆发加剧的控制机制的假说。模型分析结果表明:大型水母对中上层生物资源普遍具有显著不利影响;在大型水母、浮游动物和鲳鱼等小型中上层鱼类之间可能存在一个由大型水母爆发引发的生态系统中上层能量反馈循环;大型水母爆发初期将破坏生态系统中上层能量平衡;浮游动物生物量的波动可能是抑制大型水母爆发加剧的自然控制机制之一。

海南清澜港水母暴发期间浮游生物生态特征研究

自2010年12月-2011年5月,调查研究了海南清澜港海域的浮游生物群落特征,重点跟踪监测了2011年5月黄斑海蜇(Rhopilema hispidum)暴发期间海水营养盐水平及浮游生物群落结构变化。通过样品分析,共鉴定出浮游植物111种(变种和变型),细胞密度范围为(2.25×104)-(8.19×108)cells/L,浮游动物36种,丰度范围为19.39-25798.74ind/L,其中浮游动物以桡足类为优势种群,浮游植物以硅藻为主要优势种。在水母暴发期间浮游植物以热带骨条藻(Skeletonema tropicum)为主,浮游动物以桡足类为主。清澜港养殖区为水母水螅体提供生长发育的环境和水母暴发前期浮游植物大量增殖,为浮游动物提供充足的食物,在一定程度上促进了水母的暴发。

胶州湾海月水母(Aurelia sp．1)丰度周年变化及对浮游动物群落的影响

2009年2—12月间在胶州湾大沽河口邻近海域的逐月现场调查中,利用垂直拖网研究了海月水母碟状体和水母体的时空分布情况以及对其它浮游动物类群的影响,并探讨了海月水母的生态适应性。结果表明:海月水母的碟状体4月份开始在胶州湾出现,并且丰度逐渐增加,到6月份达到最高峰。海月水母的水母体集中在7月份大量出现。碟状体和水母体高峰期的月平均丰度分别为2.9和1.3ind/m3。碟状体开始出现时的水温平均为11.5℃,数量高峰期的水温为20.6℃,而水母体高峰期的水温为25.2℃。该海域的浮游动物总丰度(不含夜光虫)在5—7月份有个高峰期,其中5月份最高值为486.9ind/m3。7月份海月水母高峰期,浮游动物的丰度没有明显下降,两者丰度的地理变化之间也没有显著的相关性。但是通过对2006—2010年间的浮游动物各类群丰度对比,2009年海月水母暴发时夜光虫和桡足类春季丰度高值显著低于其它年份。

黄海夏季沙海蜇食物需求的估算方法及应用

自20世纪90年代末起,沙海蜇频繁暴发于东亚海域,特别是8、9月份为黄海沙海蜇生物量的高峰期/大量暴发期。研究沙海蜇的大量出现对所在生态系统的影响十分必要,最直接的影响表现在对饵料生物浮游动物的影响。本文研究了黄海2006和2007年8、9月份沙海蜇的呼吸率、摄食率,估算了沙海蜇的食物需求量及其在黄海的分布格局,获得了其每天对中、大型浮游动物现存量及生产力的潜在摄食压力。结果表明沙海蜇的食物需求量的分布格局与其生物量的分布格局一致。在沙海蜇的捕获率为最大时,2006年9月上旬沙海蜇的摄食率为47.84(0.7—215.05)mg C/(m2d)。假设中、大型浮游动物都可以作为沙海蜇的摄食对象,那么每天对中、大型浮游动物现存量及生产力的摄食压力平均分别为6.4%(0.09%—28.79%)和76.61%(1.12%—344.28%)。2006年9月下旬及2007年8月沙海蜇的食物需求比2009年9月上旬有所降低。因此,沙海蜇在暴发期间对中、大型浮游动物潜在的消耗非常大,甚至是毁灭性的。尤其是沙海蜇在高生物量站位对浮游动物的食物需求非常高,沙海蜇对浮游动物的摄食压力远超过了浮游动物本身的生产力(大于100%),这时的浮游动物远远不能满足沙海蜇的食物需求。本文的研究结果为探讨沙海蜇暴发对黄海浮游生态系统的影响程度提供了基础资料。

对黄、东海水母暴发机理的新认知

基于大量实验结果和大规模海上考察和综合分析,从基础生物学和生态学的角度,对中国近海水母暴发的机理提出一种新的理论模式:水母生活史中的大部分时间以水螅体的形式生活在海底;水母种群的暴发是水螅体对环境变异的一种应激反应,是为了逃避动荡环境、扩大分布范围、寻求新的生存空间,为种群繁衍需求更多的机会的一种生存策略。导致水母种群暴发的关键过程是海洋底层温度的变动和饵料数量的变化,全球气候变化和富营养化是中国近海水母暴发的最重要诱发因素。水母暴发是全球变化下海洋生态系统演变的一种综合体现。

秋季降温期水温和饵料量对沙蜇无性繁殖的影响

20世纪末以来,沙蜇在东亚水域频繁暴发,不仅影响正常的渔业生产活动,也给海洋生态系统健康带来极大危害。在室内模拟秋季降温条件,研究了不同水温(4种降温条件)与饵料量(0.42、1.25、和4.2μg C螅状体-1d-1)对沙蜇螅状体足囊繁殖与横裂生殖的影响。结果显示:1)秋季降温速度快慢导致的积温差异对沙蜇螅状体生成足囊数有显著影响(P<0.001),积温越大,螅状体生成的足囊数量越多,发现两者具有较好的线性关系(y=142.59x+208.39; R2=0.967),低积温条件下的足囊萌发也较少; 2)饵料量对螅状体无性繁殖的影响是有一定阈值范围的,在一定饵料量范围内,螅状体生成的足囊数量随饵料量的增加而增加,而超过一定阈值范围,过剩的饵料影响水质对沙蜇螅状体反而产生负面影响。3)秋季新繁育出来的当龄沙蜇螅状体可在越冬前发生横裂生殖,但横裂生殖的螅状体比例和放碟数量均较少。试验结果说明,全球变化背景下,积温升高、饵料量小幅增加均可能导致沙蜇秋季有性繁殖的当龄螅状体在越冬前生成更多的足囊,促进诱发翌年水母暴发。

Species-specific detection and quantification of scyphomedusae in Jiaozhou Bay,China,using a quantitative real-time PCR assay

Over the past decades,jellyfish occurred increasingly and abundantly in coastal areas worldwide.Usually,biomass of jellyfish,especially when they bloom,can be determined by visual counting.However,tiny individuals of jellyfish(e.g.,planulae,polyps,and ephyrae)are difficult to detect in the field.In this study,species-specific quantitative real-time PCR assays(qPCR)(SYBR Green I)targeting the mitochondrial 16S rDNA(mt-16S rDNA)of jellyfish were developed and were used to estimate the distribution and seasonal fluctuations of four jellyfish species(Nemopilema nomurai,Cyanea nozakii,Rhopilema esculentum,and Aurelia coerulea)in Jiaozhou Bay(JZB),China in 2013.The mt-16S rDNA of A.coerulea and N.nomurai was detected in most of the survey months and it peaked in July(1.03×10^(4) copies/L)and September(1.08×10^(6) copies/L),respectively.The mt-16S rDNA of C.nozakii occurred from August to October only with monthly mean values of 7.18-46.17 copies/L and was mainly located from the middle part to the outer part of the bay.The mt-16S rDNA of R.esculentum was the least abundant among the four species and was detected in only one sample(B2 station in March),with a value of 88.49 copies/L.The Spearman correlation test revealed that phytoplankton biomass was significantly and positively correlated with the mt-16S rDNA abundance of A.coerulea(R=0.37,P<0.01)and negatively with the mt-16S rDNA of N.nomurai(R=-0.36,P<0.01).The qPCR assay will enable the identification and quantification of jellyfish species in their whole life history and can be used as an approach in combination of the traditional jellyfish survey.

海月水母的生物学特征及其爆发

介绍了海月水母的形态、生活习性、生活史、分布、生存条件、爆发及危害等基本情况,并分析了其爆发成因,最后提出了我国海月水母及其爆发研究的工作重点:确认在我国海域爆发的海月水母究竟是外来种还是本地种,了解其种群的生活史、食性及生活习性,掌握其种群数量动态变化规律并进行数量动态监测,分析其种群数量动态与海域环境的关系,开展其爆发的危害评价工作。

Development of a real-time PCR assay(SYBR Green I) for rapid identification and quantification of scyphomedusae Aurelia sp.1 planulae

The complicated life cycle ofAurelia spp., comprising benthic asexually-reproducing polyps and sexually-reproducing medusae, makes it hard for researchers to identify and track them, especially for early stage individuals, such as planulae. To solve this problem, we developed a real-time PCR assay （SYBR Green I） to identify planulae in both cultured and natural seawater samples. Species-specific primers targeting Aurelia sp.1 mitochondrial 16S rDNA （mr 16S rDNA） regions were designed. Using a calibration curve constructed with plasmids containing the Aurelia sp. 1 mt 16S rDNA fragment and a standard curve for planulae, the absolute number of mt 16S rDNA copies per planula was determined and from that the total number ofplanulae per sample was estimated. For the field samples, a 100-fold dilution of the sample DNA combined with a final concentration of 0.2 μg/μL BSA in the PCR reaction mixture was used to remove real- time PCR inhibitors. Samples collected in Jiaozhou Bay from July to September 2012 were subsequently analyzed using this assay. Peak Aurelia sp.1 planula abundance occurred in July 2012 at stations near Hongdao Island and Qingdao offshore; abundances were very low in August and September. The real-time PCR assay （SYBR Green I） developed here negates the need for traditional microscopic identification, which is laborious and time-consuming, and can detect and quantify jellyfish planulae in field plankton samples rapidly and specifically.

海洋生态预报的研究进展与发展应用

从介绍海洋生态预报的概念和作用开始,根据赤潮、绿潮、水母暴发、珊瑚白化等海洋生态灾害的致灾种类和涉及的富营养化、低氧和致病菌等灾害指标,阐述了国内外近年来海洋生态预报的研究进展,进而探讨了河口区、陆架海以及全球大洋等不同海域差异所涉及的海洋生态预报的发展应用,论述了经验预报、统计预报和数值预报等海洋生态预报的国内外研究进展与实际应用,最后展望提出海洋生态预报在未来发展中面临的挑战和亟需解决的关键科学问题。

Experimental clearance rates of Aurelia coerulea ephyrae and medusae,and the predation impact on zooplankton in Jiaozhou Bay

The population explosion of jellyfish Aurelia coerulea occurred in Jiaozhou Bay,China in 2009.The potential predation impact of A.coerulea on zooplankton was investigated.Population clearance potential and residence time(t 1/2)for copepods were calculated from laboratory clearance rates and measurements of jellyfish size and abundance from May to August 2009 in Jiaozhou Bay.Clearance rates varied widely with prey organisms,but they were not significantly different among various prey concentrations.Medusae captured rotifers,fish larvae and hydromedusae more efficiently than fish eggs,copepods and chaetognaths.Ephyrae captured rotifers and hydromedusae more efficiently than fish larvae and copepods.Clearance rate linearly increased with the cross sectional area of A.coerulea(size from 0.3 to 7.1 cm).Water temperature also had a marked effect on clearance rate and this was related to the increased beat frequency as water temperature increased.In early May 2009,A.coerulea potentially cleared the volume of water in the Bay less than 0.001 times a day,but this value was estimated to be more than 0.3 times a day in July.The t 1/2 for copepods was less than 6 d in June and July.Abundances of copepods,hydromedusae and chaetognaths were extremely low in 2009 compared to 2008 and 2010(jellyfish non-bloom years).Large predation pressure by the A.coerulea population occurred to control zooplankton communities in Jiaozhou Bay.A.coerulea,when present at a high population level,can be a keystone species in Jiaozhou Bay and control the trophic structure here.

Natural predators of polyps of three scyphozoans:Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum

Jellyfish blooms have become a hot research topic in recent decades because they pose a serious threat to fisheries,coastal industries,tourism,and the marine ecosystem.The life cycle of scyphozoan jellyfish consists of a pelagic medusa stage and a benthic polyp stage,where asexual reproduction and strobilation of the polyps directly affect the abundance of ephyra and subsequently medusa abundance.The dynamics of polyps are affected by both environmental and biological factors,and predation by natural predators is one of the most important biological factors.Nemopilema nomurai,Aurelia coerulea,and Rhopilema esculentum are three scyphozoan species that are commonly found in Chinese coastal waters,and previous studies reported that the survivorship of polyps differs among the three species when they are exposed to the same benthic community.To identify potential natural predators of polyps of these three species in Chinese coastal waters and to determine whether the predation rates on polyps of the three species differ,we collected 39 species of macrozoobenthos from the Bohai Sea,Yellow Sea,and East China Sea from May 2014 to June 2016 and conducted predation tests and predation rate measurements.We found that the nudibranchs Pleurobranchaea novaezealandiae,Okenia plana,and Chromodoris tinctoria and the sea anemones Paracalliactis sinica,Calliactis japonica,Anthopleura incerta,and Anthopleura midori could prey on the polyps of all three scyphozoan species.The predation rates increased with the body length of the predators.The predations rates were also related to the polyp species,although the different predators showed no consistent preference for a particular species of polyp.Our results indicate that introducing predators to locations inhabited by polyps might be a way to control the benthic polyp populations and prevent subsequent jellyfish blooms.

2018年6月渤海大型水母分布特征

2018年6月使用渔业底拖网采样,对渤海大型水母进行了全面调查,调查船舶为“中渔科102”渔业科考船。本研究分析了渤海大型水母的种类组成、渔获密度与伞径大小,并对其源地进行了探讨。结果表明:本次调查共采集到海月水母、沙海蜇、海蜇、多管水母四种大型水母,其中海蜇、多管水母数量较少,各采集到一只。海月水母在渤海三湾均有分布,各海区伞径大小无显著差异且多为幼体(<10cm),密度高值区出现在渤海湾东南侧海域,可达38-221.21ind./(net·h),辽东湾海月水母出现于湾南,密度<5ind./(net·h),湾北未见;作者推测,海月水母在渤海沿岸可能存在多个源头,诸如:莱州湾与渤海湾交界近岸海域、河北近岸、辽东湾大连近岸以及北部近岸。沙海蜇在渤海分布较广,辽东湾为密度高值区,均值为(35.32±21.64)ind./(net·h),但伞径较小,均值为(12.15±6.52)cm;与此相对,渤海湾与莱州湾外侧海域沙海蜇密度虽小[<20ind./(net·h)],但伞径要显著大于辽东湾,最大伞径均值可达(33.86±7.40)cm;作者推测,沙海蜇在渤海海域发源地主要集中于辽东湾近岸,渤海湾与莱州湾,外海出现的沙海蜇可能源于辽东湾,随海流运输至此。海月水母、沙海蜇在渤海发生时间要晚于黄、东海。本研究结果可为深入分析渤海大型水母的种群动态变化、暴发机理提供基础。

盐度对海月水母幼体行为、附着和无性生殖的影响

黄渤海沿岸海参养殖池为海月水母(Aurelia aurita)幼体的重要栖息地,海参养殖池盐度波动范围极大,因此,研究不同盐度对海月水母早期生活史的影响对于理解其环境适应机制具有重要科学意义.本研究以海月水母为研究对象,探讨了不同盐度梯度(18、22、26、30、34和38)对海月水母浮浪幼虫行为、附着以及螅状幼体无性生殖的影响.结果表明,盐度对海月水母浮浪幼虫的运动行为存在显著影响,较高盐度环境(30、34和38)海月水母浮浪幼虫运动速度明显高于较低盐度环境(18、22和26);盐度对海月水母浮浪幼虫的附着有显著影响,相对较高盐度促进海月水母浮浪幼虫的附着,盐度为34时,海月水母浮浪幼虫附着率最高;海月水母螅状幼体无性生殖能适应较广的盐度范围,42d无性生殖实验周期后,18~34盐度环境下海月水母每个螅状幼体平均新繁殖螅状幼体个体数为2.8~3.7个,不同盐度处理组间无显著差异,而极高盐度(38)显著降低海月水母的出芽生殖率(1.2个).本研究表明盐度对海月水母早期生活史不同阶段幼体具有不同影响,研究结果为黄渤海沿岸海月水母环境适应机制研究提供了重要参考.