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Journal of Tropical Oceanography >

2022 , Vol. 41 >Issue 2: 142 - 148

DOI: https://doi.org/10.11978/2021071

Marine Biology

Molecular identification on the causative species jellyfish blooms in the northern South China Sea in 2019

  • DU Chong , 1, 2 ,
  • HE Jun 1 ,
  • SUN Tingting 2 ,
  • WANG Lei 2 ,
  • WANG Fanghan 2 ,
  • DONG Zhijun , 2
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  • 1. College of Life Sciences, Yantai University, Yantai 264005, China
  • 2. Muping Coastal Environment Research Station, Yantai Institute of Coastal Zone Research, Chinese Academy of Sciences, Yantai 264003, China
DONG Zhijun.

Received date: 2021-06-08

  Revised date: 2021-08-15

  Online published: 2021-08-23

Supported by

National Key Research and Development Program of China(2018YFC1406501)

Strategic Priority Research Program of the Chinese Academy of Sciences(XDA23050301)

National Natural Science Foundation of China(41876138)

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Abstract

Jellyfish blooms mostly occur in the Bohai Sea and Yellow Sea as well as in the East China Sea, but rarely in the South China Sea. We conducted morphological observations and molecular identification of blooming jellyfish in many waters of the northern South China Sea, including Haikou, Wenchang and Maoming in May 2019. Morphological results showed that the jellyfish in Haikou and Wenchang were the same species with hemispherical umbrella, pear-shaped protrusion in the lower reproductive point, and the oral arms are covered with filaments and a whip appendage at the end, which are highly similar to the Acromitus flagellatus. The jellyfish in Maoming were different: the umbrella was relatively flat hemispherical, the subgenital pits were milky protrusions and the oral arms had no filaments, which were similar to Catostylus townsendi. Based on the comparison of mitochondrial ribosomal large subunit (16S rRNA) gene sequences, the blooming jellyfish of Haikou and Wenchang were 97.5% and 97.7% similar to A. flagellatus of the eastern Pacific Ocean, and jellyfish of Maoming was 93.5% similar to C. townsendi from the Strait of Malacca. The phylogenetic tree constructed based on mitochondrial COI gene and 16S rRNA gene suggested that the jellyfish of Haikou and Wenchang were the same and belong to the same branch with A. flagellates; the jellyfish of Maoming were close to C. townsendi. Therefore, combined with morphological observations and molecular analyses, we considered Acromitus flagellatus to be the jellyfish in Haikou and Wenchang, and Catostylus sp. in Maoming.

Key words： Jellyfish bloom; South China Sea; Mitochondrial COI gene; Mitochondrial 16S rRNA gene

Cite this article

DU Chong , HE Jun , SUN Tingting , WANG Lei , WANG Fanghan , DONG Zhijun . Molecular identification on the causative species jellyfish blooms in the northern South China Sea in 2019[J]. Journal of Tropical Oceanography, 2022 , 41(2) : 142 -148 . DOI: 10.11978/2021071

近年来, 受全球气候变化和人类活动的影响, 海洋环境不断变化并出现利于水母生长繁殖的趋势, 全球多处海域出现大型水母暴发现象。水母暴发导致海洋生态系统发生变化, 对沿海渔业生产和临海工业均造成诸多负面影响(Purcell et al, 2007; Richardson et al, 2009; Purcell, 2012)。例如, 日本内海和欧洲海域水母暴发直接导致当地渔业产值大幅下降, 而在巴西沿海海域仙后水母(Cassiopea andromeda)大量入侵直接导致当地本土鱼类大量减少(Brodeur et al, 2002; Nagai, 2003; Stampar et al, 2020)。在国内, 白色霞水母(Cyanea nozakii)、海月水母(Aurelia coerulea)和沙海蜇(Nemopilema nomurai)是主要的灾害水母种类(Dong et al, 2010)。葛立军等(2004)、李建生等(2015)调查中表明, 水母暴发海域游泳动物种类和数量明显少于非暴发海域, 导致当地海域经济生物产量急剧下降, 对当地捕捞和养殖业造成了严重的经济损失。
南海位于热带气候和亚热带气候交互作用区域, 由于独特的气候特征和优越的地理位置, 因此孕育了丰富的生物多样性(李纯厚 等, 2005; 陈清潮, 2011)。南海大型水母物种丰富, 常见大型水母主要为马来沙水母(Sanderia malayensis)、白色霞水母、紫色霞水母(Cyanea purpurea)、陈嘉庚水母(Acromitus tankahkeei)、鞭腕水母(Acromitus flagellatus)和黄斑海蜇(Rhopilema hispidum)等(陈小银, 2011; 岑竞仪 等, 2012)。水母暴发多发生在黄渤海和东海海域, 在南海海域较为少见(Dong et al, 2010)。其中, 岑竞仪等(2012)曾报道在海南文昌海域出现水母暴发现象, 黄斑海蜇作为原因种最大密度可达30只·m-3。2019年5月在海南海口和广东茂名附近海域均出现了水母暴发现象(聂元剑, 2019), 但原因种未知。
目前, 分子生物技术的快速发展为生物物种的快速鉴定提供了新思路, 并广泛用于水母物种的鉴定。例如, 张姝等(2009)利用线粒体COI和16S rRNA基因序列对我国海区的海蜇(Rhopilema esculentum)和沙海蜇进行了分子鉴定, 发现黄海海域的沙海蜇与日本的越前水母(Nemopilema nomurai)为同一物种; Mutlu等(2020)基于COI基因对地中海未知水母进行物种鉴定, 并将该水母命名为Chrysaora pseudoocellata sp. nov.; Seo等(2021)基于线粒体COI和16S rRNA基因序列对海月水母水螅体的遗传多样性进行研究, 结果表明韩国海月水母(A. coerulea)的不同地理群体存在明显的遗传分化。因此, 本研究利用分子系统学鉴定技术, 结合形态分类学对2019年南海北部近海暴发水母原因种进行鉴定, 摸清南海海域水母暴发的物种种类, 为深入研究南海海域灾害水母的暴发机制提供了基础信息。

1 材料与方法

1.1 样品采集

2019年6月分别在海口(20°2′37″N, 110°19′18″E)、文昌(19°34′11″N, 110°49′59″E)和茂名(21°23′35″N, 111°11′56″E)附近海域采集暴发水母, 每个海域各采集1个水母个体(图1)。对暴发水母样品形态学鉴定和拍照记录后, 取伞部部分组织保存在95%的乙醇中, -20℃保存。
图1 南海采样地点分布图

该图基于国家测绘地理信息局标准地图服务网站下载的审图号为GS(2020)4632的标准地图制作, 底图无修改

Fig. 1 Map of sampling sites in the South China Sea.

The map is based on the standard map downloaded from the standard map service website of the National Bureau of Surveying, Mapping and Geographic Information with the review number GS(2020)4632

1.2 形态学鉴定方法

参考《中国动物志》(高尚武 等, 2002)和《World Atlas of Jellyfish》(Jarms et al, 2019)中钵水母形态与结构的描述, 对采集水母样品的伞部形状大小、缘瓣形状和数目、感受器位置数目、生殖下腔、口腕及附属物和整体特征等进行观察记录。

1.3 基因组DNA提取、扩增及测序

取出少量样品伞部组织, 用双蒸水反复冲洗, 采用海洋动物组织基因组DNA提取试剂盒(天根, 北京天根生化科技有限公司)对水母样品DNA进行提取, 基因组DNA保存在-20℃的洗脱缓冲液中。
线粒体COI基因片段扩增采用通用引物LCO1490: 5′-GGTCAACAAATCATAAAGATATT GG-3′; HCO2198: 5′-TAAACTTCAGGGTGACCAAAAAATCA-3′ (Folmer et al, 1994)。反应体系为25μL, 包括12.5μL 2×Taq PCR StarMix, 10μmol·L-1引物各1μL, 3μL DNA模板, 其他以灭菌超纯水补足体系。扩增程序为: 94℃预变性3min, 然后35个循环(包括94℃变性30s, 54℃退火30s), 循环结束后72℃延伸7min, 4℃低温保存。线粒体16S rRNA基因片段扩增采用引物16S-L: 5′-GACTGTTTACCA ACATA-3′; 16S-H: 5′-CATAATTCAACATCGAGGT- 3′ (Ender et al, 2003)。反应体系为25μL, 包括12.5μL 2×Taq PCR StarMix, 10μmol·L-1引物各1μL, 3μL DNA模板, 其他以灭菌超纯水补足体系。16S rRNA基因PCR扩增程序为: 先预处理5个循环(94℃预变性1min, 45℃退火50s, 72℃延伸1min), 然后30个循环(包括94℃变性50s, 50℃退火1min, 72℃延伸1min), 循环结束后72℃延伸5min, 4℃低温保存。PCR扩增在Takara TP350 PCR仪上进行。
取PCR扩增产物5μL进行1%琼脂糖凝胶电泳, 电泳程序为: 电压110V运行30min, 在凝胶电泳成像仪器系统拍照记录检测PCR扩增结果。将凝胶系统检测含有扩增片段的样品送上海生工生物工程技术服务有限公司进行纯化和测序, 测序使用ABI测序仪(3730xl DNA Analyer)双向测序。

1.4 数据分析

用软件BioEdit对DNA序列进行人工校正和编辑, 用MEGA7软件对测定的样品序列进行碱基含量分析。在GenBank中收集根口水母目(Rhizostomeae)的COI基因序列和16S rRNA基因序列, 每个科选择一个代表种, 共收集到11个COI基因序列和18个16S rRNA基因序列。采用邻接法(Neighbor-joining)构建系统发育树, 用Bootstrap 1000检验分子系统树各分支置信度, 遗传距离采用Kimura双参数模型计算。

2 结果

2.1 形态学结果

海口和文昌附近海域暴发水母形态相同(图2a), 主要形态特征为: 水母伞部呈半球状, 伞径约130mm, 外伞部表面有细小颗粒。伞缘8个感觉器, 每1/8伞缘有4对长舌状缘瓣。生殖下穴呈梨形突起, 彼此相通。口腕8条, 为3翼型, 约与伞径等长, 布满短丝状物, 末端有1条鞭状附属物。生活时伞部有黑褐色斑点, 口腕乳白色。初步形态学鉴定, 该水母形态与鞭腕水母相似(高尚武 等, 2002; 洪惠馨, 2014; Jarms et al, 2019)。
图2 南海北部近海暴发水母形态图

a. 海口和文昌附近海域暴发水母; b. 茂名附近海域暴发水母。um表示水母伞部, fa表示鞭状附属物, oa表示水母口腕, ml表示缘瓣

Fig. 2 Morphological character of jellyfish in the South China Sea.

Jellyfish collected in Haikou and Wenchang offshore area (a), and in Maoming offshore area (b). um: umbrella; fa: flagelliform appendage; oa: oral arms; ml: marginal lappets

茂名附近海域暴发水母(图2b)不同于海口和文昌海域暴发水母, 主要形态特征为: 伞部呈扁平半球状, 伞径约120mm, 外伞部表面光滑有颗粒状突起, 胶质层坚硬。伞缘8个感觉器, 缘瓣数目较多, 缘瓣之间的裂缝较深。生殖下穴呈乳状突起, 彼此相通。口腕8条, 为3翼型, 比伞部直径略长, 末端逐渐变钝, 没有附属物。生活时整个个体呈白色, 伞部布满深棕色斑点。初步形态学鉴定, 该水母形态与端棍水母(Catostylus townsendi)相似(洪惠馨, 2014; Jarms et al, 2019)。

2.2 分子鉴定结果

海口、文昌和茂名附近海域水母样品COI基因序列扩增测序长度为658bp, 上传至GeneBank获得序列号MZ028018~MZ028020。海口和文昌附近海域暴发水母样品16S rRNA基因序列扩增测序长度为527bp, 而茂名海域暴发水母16S rRNA基因长度为522bp, 上传至GeneBank获得序列号MZ031038~ MZ031040。暴发水母COI和16S rRNA基因片段碱基组成显示, 所有基因片段的A+T含量均明显高于C+G含量, 表现出明显的A/T偏好, 详细信息见表1。
表1 各地区暴发水母碱基组成及GenBank序列号

Tab. 1 Base composition and GenBank sequence number of jellyfish in different offshore areas

采样地点 A/% T/% C/% G/% 总长度/bp GenBank序列号
COI基因序列 海口 31.5 28.3 17.9 22.3 658 MZ028018
文昌 31.3 28.3 17.9 22.5 658 MZ028019
茂名 35.4 27.5 18.1 19.0 658 MZ028020
16S rRNA序列 海口 26.4 38.9 17.8 16.9 527 MZ031038
文昌 26.2 38.9 17.8 17.1 527 MZ031039
茂名 38.1 29.9 14.9 17.0 522 MZ031040
利用NCBI中的BLAST (http://www.ncbi.nlm.gov/BLAST/)检索, 与GenBank中已记录水母COI和16S rRNA基因序列进行相似性比对。结果显示, COI基因序列对比中, 海口和文昌两地附近海域暴发水母序列相似性高(>99.0%), 与印尼海域鞭腕水母序列(序列号: MN395685)相似性为89.9%和89.6%, 而茂名附近海域暴发水母序列与印尼海域端棍水母序列(序列号: MN395693)相似性为85.9%。在16S rRNA基因序列对比中, 海口和文昌两地海域暴发水母序列相似性高(>99.0%), 与东太平洋的鞭腕水母序列(序列号: KY610576)相似性达到97.5%和97.7%, 茂名附近海域暴发水母序列与马六甲海峡的端棍水母序列(序列号: KY610587)相似性为93.5%。
基于COI基因序列系统发育树显示, 海口和文昌附近海域暴发水母首先聚为一支, 然后和鞭腕水母汇聚成一支, 自展支持率均为100。而茂名附近海域暴发水母与端棍水母聚在一起, 自展支持率为74 (图3)。
图3 基于COI基因序列的根口水母目邻接系统发育树

图中HK、WC和MM分别代表海口、文昌和茂名附近海域暴发水母, 各支上数据为Bootstrap 1000给出自展支持值, 只显示大于50的自展值

Fig. 3 Neighbor joining phylogenetic tree built based on the COI sequences of Rhizostomeae.

HK, WC and MM represent jellyfish in Haikou, Wenchang and Maoming offshore areas, respectively. The number on each branch is Bootstrap 1000 to give bootstrap values (only the one larger than 50 is shown)

基于16S rRNA基因序列系统发育树显示, 海口和文昌附近海域暴发水母先聚为一支, 又与鞭腕水母聚为一支, 自展支持率均为100。茂名海域暴发水母与端棍水母聚为一支, 自展支持率为97 (图4)。海口和文昌附近海域水母的遗传距离是0.006, 与东太平洋鞭腕水母的遗传距离分别为0.025和0.023。而茂名附近海域水母与马六甲海峡端棍水母遗传距离为0.048。
图4 基于16S rRNA基因序列根口水母目邻接系统发育树

图中HK、WC和MM分别代表海口、文昌和茂名附近海域暴发水母, 各支上数据为Bootstrap 1000给出自展支持值, 只显示大于50的自展值

Fig. 4 Neighbor joining phylogenetic tree based on the 16S rRNA sequences of Rhizostomeae.

HK, WC and MM represent jellyfish in Haikou, Wenchang and Maoming offshore areas, respectively. The number on each branch is Bootstrap 1000 to give bootstrap values (only the one larger than 50 is shown)

3 讨论

水母特征通常会受到生境变化的影响, 环境的多样性和复杂性可能使水母的形态特征发生微小的变化, 从而影响水母的鉴别和分类。所以从传统的基于几个最显著的形态特征来鉴别物种的方法, 并不能有效解决现有物种鉴定的问题(Dawson et al, 2001; 郑连明 等, 2008)。近年来分子分类学快速发展, 相比于核DNA, 线粒体基因由于具有结构简单、进化速率高和母系遗传等特点, 适合物种进化分析研究(Brown et al, 1979)。线粒体中的COI基因和16S rRNA基因序列具有高效和成本低的优点, 越来越多地被用于水母的分类研究(郑连明 等, 2008; 张姝 等, 2009; 王建艳 等, 2013; 张珰妮 等, 2015)。本研究利用分子生物学技术, 对南海北部近海海域的暴发水母线粒体COI基因序列和线粒体16S rRNA基因序列进行了测定。通过分析线粒体基因序列信息, 构建系统发育树和计算遗传距离, 为形态学观察结果提供分子水平的分类依据。
形态学观察显示, 海口和文昌附近海域暴发种为同一种水母, 该物种具有伞部表面有细小颗粒和黑褐色的斑点、生殖下穴呈梨形突起、口腕布满短丝状物和末端有鞭状附属物等明显特征, 初步判定为鞭腕水母。鞭腕水母属于刺胞动物门(Cnidaria), 钵水母纲(Scyphozoa), 根口水母目(Rhizostomeae), 鞭棍水母科(Catostylidae), 端鞭水母属(Acromitus)。鞭腕水母内环网状管和感觉辐管相连, 典型特征是口腕布满短丝状物并在末端有1条鞭状丝状附属物, 主要分布在中国南海海域(高尚武 等, 2002; 洪惠馨, 2014; Jarms et al, 2019)。在我国南海近海海域有两种端鞭水母, 陈嘉庚水母和鞭腕水母(高尚武 等, 2002; 洪惠馨, 2014)。海口和文昌海域水母不同于陈嘉庚水母, 具有个体大(伞径约130mm)和口腕长度略与伞径等长的特征。因此, 初步判定海口和文昌海域水母为鞭腕水母。而茂名附近海域暴发种具有外伞部表面光滑布满褐色斑点、伞缘缘瓣数目较多裂缝较深、生殖下穴呈乳状突起、口腕8条没有附属物等特征, 与端棍水母形态相似。端棍水母属于刺胞动物门(Cnidaria), 钵水母纲(Scyphozoa), 根口水母目(Rhizostomeae), 鞭棍水母科(Catostylidae), 端棍水母属(Catostylus)。端棍水母内网络系统宽网眼大, 内网络直接与感觉器和间辐管连接, 外网延伸到缘瓣, 网眼小, 口腕没有附属物(洪惠馨, 2014; Jarms et al, 2019)。
基于线粒体基因碱基组成显示, 两种水母两种线粒体基因均表现出较高的A+T的含量, 符合水母类线粒体基因组A+T含量普遍偏高的特点(程方平 等, 2012; Dong et al, 2015; 刘青青 等, 2018)。基于线粒体COI基因序列和线粒体16S rRNA基因序列构建的系统发育树结果显示, 海口、文昌和茂名附近海域暴发水母基因序列明显分为两支, 海口和文昌的水母先聚在一起后与鞭腕水母聚到一支, 说明海口和文昌海域水母与鞭腕水母关系十分接近。而茂名附近海域水母和端棍水母聚为一支, 说明茂名附近海域水母与端棍水母有较近的亲缘关系。基于线粒体16S rRNA基因序列的遗传距离显示, 海口和文昌附近海域暴发水母间的遗传距离为0.006, 基因序列差异很低属于种内遗传差异, 可以认为海口和文昌附近海域暴发水母为同一种。而海口和文昌海域水母与东太平洋鞭腕水母遗传距离为0.025和0.023, 茂名海域水母与马六甲海峡端棍水母遗传距离是0.048。基于线粒体16S rRNA基因的研究结果显示水螅水母的种内遗传距离为0~0.047, 而立方水母和钵水母的种内遗传距离在0~0.05之间(张珰妮 等, 2015; 刘瑞娟 等, 2016)。因此, 本研究中海口和文昌海域暴发水母与东太平洋鞭腕水母间的遗传分化可以认为是种内遗传分化。而茂名海域暴发水母与马六甲海峡端棍水母遗传分化程度较大, 推测茂名海域暴发水母可能为端棍水母隐存种。本研究还揭示水母线粒体16S rRNA基因比线粒体COI基因更保守, 适合水母分子系统分类的研究, 这和何劲儒(2014)的研究结论一致。
结合传统的形态学观察结果, 本研究从分子水平上证实海口和文昌附近海域暴发水母为鞭腕水母(Acromitus flagellatus), 而茂名附近海域暴发水母为端棍水母(Catostylus sp.)。本研究鉴定了2019年5月南海多地出现水母暴发的原因种, 摸清南海北部海域暴发水母物种种类, 为深入研究南海海域灾害水母的暴发机制提供了基础信息。

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