Study on the tempo-spatial distribution of green macroalgal micro-propagules along the coasts of Jiangsu and Shandong provinces
-
摘要: 为研究暴发浒苔绿潮的黄海近岸水体中大型绿藻微观繁殖体时空分布和种群演替，以及绿潮对海域繁殖体群落结构的影响，本研究在江苏－山东沿岸设置4个站位，开展周年取样调查研究。通过室内培养和分子检测结合的方法，共鉴定出6种大型绿藻：缘管浒苔(Ulva linza)、浒苔(U. prolifera)、扁浒苔(U. compressa)、孔石莼(U. pertusa)、未命名种(Ulva sp.) 和盘苔(Blidingia sp.)。定量研究结果表明，各站位微观繁殖体丰度和种类组成具有明显时空变化特征，苏北浅滩与青岛近岸周年绿藻微观繁殖体丰度明显高于连云港燕尾港和日照岚山港站位；各站位丰度呈双峰型波动，春季到初夏达最高，秋季为次高峰，其他季节丰度较低。群落结构组成研究表明，苏北浅滩周年存在较高丰度的浒苔微观繁殖体(平均为77株/L)，其中造成黄海大规模绿潮的“漂浮生态型”浒苔占绝对优势，年均占浒苔微观繁殖体总数的75.7%；青岛近岸微观繁殖体种类组成波动显著，在夏季大规模浒苔绿潮到达时，优势种从缘管浒苔转变成浒苔，其中，“漂浮生态型”浒苔微观繁殖体数量急剧上升，并在绿潮消退时快速消失。对比研究显示，苏北浅滩海域是浒苔微观繁殖体“栖息地”，对维持浒苔种群、造成绿潮连年暴发具有重要作用；青岛近岸微观繁殖体群落受到大规模漂浮浒苔的季节性扰动，其组成结构在绿潮暴发期发生显著变化。Abstract: To study the tempo-spatial distribution and species succession of green macroalgal micro-propagules in the coastal waters affected by the green tides in the Yellow Sea, a year-round survey was conducted at the four stations (ZGS, YWG, LSG and YY) along the coasts of Jiangsu and Shandong provinces. Through the lab-culture and molecular analysis, there were six green macroalgal species detected, including 4 ulvoid species (Ulva linza, U. prolifera, U. compressa and U. pertusa), an unclassified Ulva sp. and one Blidingia sp. Quantitative analyses revealed evident tempo-spatial variations of the abundance and species composition of micro-propagules at the four stations. The abundances of micro-propagules were significantly higher at ZGS and YY than those at YWG and LSG. A seasonal bi-peak pattern of the abundances was consistent throughout the four stations. The numbers of micro-propagules were generally highest during the spring to early summer, and reached a minor peak in fall, but were low in other seasons. The U. prolifera propagules (77 individuals/L in average over the year) were existing at ZGS of the Subei Shoal throughout the year, and majority (75.7% in average) of them was the ‘floating’ type, an ecotype causing the green tides in the Yellow Sea. In comparison, species composition of micro-propagules fluctuated significantly at YY of Qingdao. The dominant species switched from U. linza to U. prolifera in summer when the large-scale floating biomass of U. prolifera approached the coasts of Qingdao. The quantity of ‘floating’ type of U. prolifera increased dramatically in this period and descended rapidly with the cessation of the green tide. This study suggested that the Subei Shoal is an important ‘habitat’ for U. prolifera, which plays a crucial role in maintaining the U. prolifera population and causing the successive green tides. Whereas in Qingdao coastal water, the large-scale floating U. prolifera biomass could temporarily influence the local micro-propagule community, and cause a rapid shift of the micro-propagule species during the blooming season of the green tides.
-
Key words:
- Ulva prolifera /
- green tides /
- micro-propagules /
- species succession
-
图 1 江苏−山东沿岸4个取样站位
Fig. 1 Four sampling locations along the coasts of Jiangsu and Shandong Provinces
图 2 2016年3月至2017年3月江苏−山东沿岸各站位绿藻微观繁殖体数量变化（平均值±标准误差，*代表未采样）
Fig. 2 Micro-propagule abundance at four sampling locations along the coasts of Jiangsu and Shandong provinces from March 20016 to March 2017 (mean±SE, * indicates no samples collected)
图 3 2016年3月至2017年3月ZGS和YY站绿藻微观繁殖体种类组成（*代表未采样）
Fig. 3 Species composition of the mciro-propagule communities at ZGS and YY stations from March 2016 to March 2017 (* indicates no samples collected)
图 4 2016年3月至2017年3月ZGS、YY站浒苔微观繁殖体丰度及“漂浮生态型”浒苔占比（*代表未采样）
Fig. 4 Abundance of the total and ‘floating’ type of U. prolifera at ZGS and YY stations from March 2016 to March 2017 (* indicates no sample collected)
-
[1] Merceron M, Antoine V, Auby I, et al. In situ growth potential of the subtidal part of green tide forming Ulva spp. stocks[J]. Science of the Total Environment, 2007, 384(1/3): 293−305. [2] Charlier R H, Morand P, Finkl C W, et al. Green tides on the Brittany coasts[C]//Proceedings of 2006 IEEE US/EU Baltic International Symposium. Klaipeda, Lithuania: IEEE, 2006. [3] Nelson T A, Haberlin K, Nelson A V, et al. Ecological and physiological controls of species composition in green macroalgal blooms[J]. Ecology, 2008, 89(5): 1287−1298. doi: 10.1890/07-0494.1 [4] Shimada S, Hiraoka M, Nabata S, et al. Molecular phylogenetic analyses of the Japanese Ulva and Enteromorpha (Ulvales, Ulvophyceae), with special reference to the free-floating Ulva[J]. Phycological Research, 2003, 51(2): 99−108. doi: 10.1111/j.1440-1835.2003.tb00176.x [5] Sun Song, Wang Fan, Li Chaolun, et al. Emerging challenges: massive green algae blooms in the yellow sea[J/OL]. Nature Precedings, (2008-09-10). http://precedings.nature.com/documents/2266/version/1. [6] Liu Dongyan, Keesing J K, Xing Qianguo, et al. World's largest macroalgal bloom caused by expansion of seaweed aquaculture in China[J]. Marine Pollution Bulletin, 2009, 58(6): 888−895. doi: 10.1016/j.marpolbul.2009.01.013 [7] Wang Zongling, Xiao Jie, Fan Shiliang, et al. Who made the world's largest green tide in China? - an integrated study on the initiation and early development of the green tide in Yellow Sea[J]. Limnology and Oceanography, 2015, 60(4): 1105−1117. doi: 10.1002/lno.10083 [8] Hu Chuanmin, Li Daqiu, Chen Changsheng, et al. On the recurrent Ulva prolifera blooms in the Yellow Sea and East China Sea[J]. Journal of Geophysical Research: Oceans, 2010, 115(C5): C05017. [9] 叶乃好, 张晓雯, 毛玉泽, 等. 黄海绿潮浒苔(Enteromorpha prolifera)生活史的初步研究[J]. 中国水产科学, 2008, 15(5): 853−859.Ye Naihao, Zhang Xiaowen, Mao Yuze, et al. Life history of Enteromorpha prolifera under laboratory conditions[J]. Journal of Fishery Sciences of China, 2008, 15(5): 853−859. [10] 李瑞香, 吴晓文, 韦钦胜, 等. 不同营养盐条件下浒苔的生长[J]. 海洋科学进展, 2009, 27(2): 211−216.Li Ruixiang, Wu Xiaowen, Wei Qinsheng, et al. Growth of Enteromorpha prolifera under different nutrient conditions[J]. Advances in Marine Science, 2009, 27(2): 211−216. [11] 范士亮, 傅明珠, 李艳, 等. 2009-2010年黄海绿潮起源与发生过程调查研究[J]. 海洋学报, 2012, 34(6): 187−194.Fan Shiliang, Fu Mingzhu, Li Yan, et al. Origin and development of Huanghai (Yellow) Sea green-tides in 2009 and 2010[J]. Haiyang Xuebao, 2012, 34(6): 187−194. [12] 乔方利, 王关锁, 吕新刚, 等. 2008与2010年黄海浒苔漂移输运特征对比[J]. 科学通报, 2011, 56(21): 2236−2242.Qiao Fangli, Wang Guansuo, Lü Xingang, et al. Drift characteristics of green macroalgae in the Yellow Sea in 2008 and 2010[J]. Chinese Science Bulletin, 2011, 56(21): 2236−2242. [13] 于仁成, 孙松, 颜天, 等. 黄海绿潮研究: 回顾与展望[J]. 海洋与湖沼, 2018, 49(5): 942−949.Yu Rencheng, Sun Song, Yan Tian, et al. Progresses and perspectives on green-tide studies in the Yellow Sea[J]. Oceanologia et Limnologia Sinica, 2018, 49(5): 942−949. [14] 王宗灵, 傅明珠, 肖洁, 等. 黄海浒苔绿潮研究进展[J]. 海洋学报, 2018, 40(2): 1−13.Wang Zongling, Fu Mingzhu, Xiao Jie, et al. Progress on the study of the Yellow Sea green tides caused by Ulva prolifera[J]. Haiyang Xuebao, 2018, 40(2): 1−13. [15] Lotze H K, Worm B, Sommer U. Strong bottom-up and top-down control of early life stages of macroalgae[J]. Limnology and Oceanography, 2001, 46(4): 749−757. doi: 10.4319/lo.2001.46.4.0749 [16] Hoffmann M A, Santelices G B. Banks of algal microscopic forms: hypotheses on their functioning and comparisons with seed banks[J]. Marine Ecology Progress Series, 1991, 79: 185−194. doi: 10.3354/meps079185 [17] Lotze H K, Schramm W, Worm S B. Control of macroalgal blooms at early developmental stages: Pilayella littoralis versus Enteromorpha spp.[J]. Acta Oecologia, 1999, 119(1): 46−54. doi: 10.1007/s004420050759 [18] 黄娟, 吴玲娟, 高松, 等. 黄海绿潮分布年际变化分析[J]. 激光生物学报, 2014, 23(6): 572−578.Huang Juan, Wu Lingjuan, Gao Song, et al. Analysis on the interannual distribution variation of green tide in Yellow Sea[J]. Acta Laser Biology Sinica, 2014, 23(6): 572−578. [19] Bao Min, Guan Weibing, Yang Yang, et al. Drifting trajectories of green algae in the western Yellow Sea during the spring and summer of 2012[J]. Estuarine, Coastal and Shelf Science, 2015, 163: 9−16. doi: 10.1016/j.ecss.2015.02.009 [20] Huo Yuanzi, Han Hongbin, Hua Liang, et al. Tracing the origin of green macroalgal blooms based on the large scale spatio-temporal distribution of Ulva microscopic propagules and settled mature Ulva vegetative thalli in coastal regions of the Yellow Sea, China[J]. Harmful Algae, 2016, 59: 91−99. doi: 10.1016/j.hal.2016.09.005 [21] Song Wei, Li Yan, Fang Song, et al. Temporal and spatial distributions of green algae micro-propagules in the coastal waters of the Subei Shoal, China[J]. Estuarine, Coastal and Shelf Science, 2014, 163: 29−35. [22] Liu Dongyan, Keesing J K, He Peimin, et al. The world's largest macroalgal bloom in the Yellow Sea, China: Formation and implications[J]. Estuarine, Coastal and Shelf Science, 2013, 129: 2−10. doi: 10.1016/j.ecss.2013.05.021 [23] 方松, 王宗灵, 李艳, 等. 南黄海及长江口附近海域绿潮暴发前期微观繁殖体的动态变化[J]. 海洋学报, 2012, 34(4): 147−154.Fang Song, Wang Zongling, Li Yan, et al. The dynamics of micro-propagules before the green tide (Ulva prolifera) outbreak in the Southern Huanghai Sea and Changjiang (Yangtze) River Estuary area[J]. Haiyang Xuebao, 2012, 34(4): 147−154. [24] Li Yan, Song Wei, Xiao Jie, et al. Tempo-spatial distribution and species diversity of green algae micro-propagules in the Yellow Sea during the large-scale green tide development[J]. Harmful Algae, 2014, 39: 40−47. doi: 10.1016/j.hal.2014.05.013 [25] Zhang Jianheng, Zhao Peng, Huo Yuanzi, et al. The fast expansion of Pyropia aquaculture in “Sansha” regions should be mainly responsible for the Ulva blooms in Yellow Sea[J]. Estuarine, Coastal and Shelf Science, 2017, 189: 58−65. doi: 10.1016/j.ecss.2017.03.011 [26] Berges J A, Franklin D J, Harrison P J. Evolution of an artificial seawater medium: improvements in enriched seawater, artificial water over the last two decades[J]. Journal of Phycology, 2001, 37(6): 1138−1145. doi: 10.1046/j.1529-8817.2001.01052.x [27] Hwangbo K, Son S H, Lee J S, et al. Rapid and simple method for DNA extraction from plant and algal species suitable for PCR amplification using a chelating resin Chelex 100[J]. Plant Biotechnology Reports, 2010, 4(1): 49−52. doi: 10.1007/s11816-009-0117-4 [28] Xiao Jie, Li Yan, Song Wei, et al. Discrimination of the common macroalgae (Ulva and Blidingia) in coastal waters of Yellow Sea, northern China, based on restriction fragment-length polymorphism (RFLP) analysis[J]. Harmful Algae, 2013, 27(7): 130−137. [29] Leskinen E, Pamilo P. Evolution of the ITS Sequences of ribosomal DNA in Enteromorpha (Chlorophyceae)[J]. Hereditas, 1997, 126(1): 17−23. [30] Hayden H S, Blomster J, Maggs C A, et al. Linnaeus was right all along: Ulva and Enteromorpha are not distinct genera[J]. European Journal of Phycology, 2003, 38(3): 277−294. doi: 10.1080/1364253031000136321 [31] Shimada, S, Yokoyama, N, Ar ai, S, et al. Phylogeography of the genus Ulva (Ulvophyceae, Chlorophyta), with special reference to the Japanese freshwater and brackish taxa[J]. Journal of Applied Phycology, 2008, 20(5): 979−989. doi: 10.1007/s10811-007-9296-y [32] Zhao Jin, Jiang Peng, Qin Song, et al. Genetic analyses of floating Ulva prolifera in the Yellow Sea suggest a unique ecotype[J]. Estuarine, Coastal and Shelf Science, 2015, 163: 96−102. doi: 10.1016/j.ecss.2015.05.027 [33] 田晓玲, 霍元子, 陈丽平, 等. 江苏如东近海绿潮藻分子检测与类群演替分析[J]. 科学通报, 2011, 56(4/5): 309−317.Tian Xiaoling, Huo Yuanzi, Chen Liping, et al. Molecular detection and analysis of green seaweeds from Rudong coasts in Jiangsu Province[J]. Chinese Science Bulletin, 2011, 56(4/5): 309−317. [34] Duan Weijin, Guo Lixin, Dong Sun, et al. Morphological and molecular characterization of free-floating and attached green macroalgae Ulva spp. in the Yellow Sea of China[J]. Journal of Applied Phycology, 2012, 24(1): 97−108. doi: 10.1007/s10811-011-9654-7 [35] Zhao Jin, Jiang Peng, Qiu Ri, et al. The Yellow Sea green tide: a risk of macroalgae invasion[J]. Harmful Algae, 2018, 77: 11−17. doi: 10.1016/j.hal.2018.05.007 [36] Taylor R, Fletcher R L, Raven J A. Preliminary studies on the growth of selected ‘green tide’ algae in laboratory culture: effects of irradiance, temperature, salinity and nutrients on growth rate[J]. Botanica Marina, 2005, 44(4): 327−336. [37] 刘峰, 刘兴凤, 金柘, 等. 苏北浅滩沉积物中的大型绿藻微观繁殖体的垂直分布和物种多样性[J]. 海洋与湖沼, 2018, 49(5): 983−990.Liu Feng, Liu Xingfeng, Jin Zhe, et al. Vertical distribution and species diversity of microscopic propagules of green macroalgae in sediments of Subei Shoal, China[J]. Oceanologia et Limnologia Sinica, 2018, 49(5): 983−990. [38] 吴晓文, 李瑞香, 徐宗军, 等. 营养盐对浒苔生长影响的围隔生态实验[J]. 海洋科学进展, 2010, 28(4): 538−544.Wu Xiaowen, Li Ruixiang, Xu Zongjun, et al. Mesocosm experiments of nutrient effects on Enteromorpha prolifera growth[J]. Advances in Marine Science, 2010, 28(4): 538−544. [39] Li Youxun, Zhang Xiaowen, Xu Dong, et al. Differential gene expression in Ulva prolifera under low light and low temperature conditions[J]. Current Genetics, 2012, 58(4): 235−244. doi: 10.1007/s00294-012-0380-8 [40] Lin Apeng, Wang Chao, Pan Guanghua, et al. Diluted seawater promoted the green tide of Ulva prolifera (Chlorophyta, Ulvales)[J]. Phycological Research, 2011, 59(4): 295−304. doi: 10.1111/j.1440-1835.2011.00629.x [41] Luo Minbo, Liu Feng, Xu Zhaoli. Growth and nutrient uptake capacity of two co-occurring species, Ulva prolifera and Ulva linza[J]. Aquatic Botany, 2012, 100: 18−24. doi: 10.1016/j.aquabot.2012.03.006 [42] Wang Ying, Wang You, Zhu Lin, et al. Comparative studies on the ecophysiological differences of two green tide macroalgae under controlled laboratory conditions[J]. PLoS One, 2012, 7(8): e38245. doi: 10.1371/journal.pone.0038245 [43] Xiao Jie, Zhang Xiaohong, Gao Chunlei, et al. Effect of temperature, salinity and irradiance on growth and photosynthesis of Ulva prolifera[J]. Acta Oceanologica Sinica, 2016, 35(10): 114−121. doi: 10.1007/s13131-016-0891-0 [44] Liu Feng, Pang Shaojun, Zhao Xiaobo, et al. Quantitative, molecular and growth analyses of Ulva microscopic propagules in the coastal sediment of Jiangsu province where green tides initially occurred[J]. Marine Environmental Research, 2012, 74: 56−63. doi: 10.1016/j.marenvres.2011.12.004 [45] Liu Feng, Pang Shaojun, Chopin T, et al. Understanding the recurrent large-scale green tide in the Yellow Sea: Temporal and spatial correlations between multiple geographical, aquacultural and biological factors[J]. Marine Environmental Research, 2013, 83: 38−47. doi: 10.1016/j.marenvres.2012.10.007 [46] 姜鹏, 赵瑾. 黄海绿潮浒苔漂浮生态型的发现与启示[J]. 海洋与湖沼, 2018, 49(5): 959−966.Jiang Peng, Zhao Jin. Identification of floating ecotype of Ulva prolifera: implication for the green tide in the Yellow Sea[J]. Oceanologia et Limnologia Sinica, 2018, 49(5): 959−966. [47] Zhang Qingchun, Yu Rencheng, Chen Zhenfan, et al. Genetic evidence in tracking the origin of Ulva prolifera blooms in the Yellow Sea, China[J]. Harmful Algae, 2018, 78: 86−94. doi: 10.1016/j.hal.2018.08.002 [48] 徐福祥, 高志强, 郑翔宇, 等. 基于MODIS数据的2016年黄海绿潮灾害动态监测研究[J]. 海洋科学, 2017, 41(5): 80−84.Xu Fuxiang, Gao Zhiqiang, Zheng Xiangyu, et al. Spatial and temporal distribution characteristic of green tides in the Yellow Sea in 2016 based on MODIS data[J]. Marine Sciences, 2017, 41(5): 80−84. [49] Geng Huixia, Yan Tian, Zhou Mingjiang, et al. Comparative study of the germination of Ulva prolifera gametes on various substrates[J]. Estuarine, Coastal and Shelf Science, 2015, 163: 89−95. doi: 10.1016/j.ecss.2014.12.026 [50] Shi Xiaoyong, Qi Mingyan, Tang Hongjie, et al. Spatial and temporal nutrient variations in the Yellow Sea and their effects on Ulva prolifera blooms[J]. Estuarine, Coastal and Shelf Science, 2015, 163: 36−43. doi: 10.1016/j.ecss.2015.02.007 [51] Lin Hanzhi, Jiang Peng, Zhang Jiaxu, et al. Genetic and marine cyclonic eddy analyses on the largest macroalgal bloom in the world[J]. Environmental Science & Technology, 2011, 45(14): 5996−6002. [52] 谭林涛, 徐雯雯, 郑富强, 等. 山东紫菜“热”的“冷”思考[J]. 中国水产, 2018(7): 46−48.Tan Lintao, Xu Wenwen, Zheng Fuqiang, et al. Thoughts on Pyropia farming in Shandong[J]. China Fisheries, 2018(7): 46−48. [53] 李欣钰, 康美华, 侯鹏飞, 等. 2013年胶州湾溶解营养盐时空分布特征[J]. 海洋环境科学, 2016, 35(3): 334−342.Li Xinyu, Kang Meihua, Hou Pengfei, et al. Spatial and temporal distribution of nutrient in Jiaozhou Bay in 2013[J]. Marine Environmental Science, 2016, 35(3): 334−342. [54] Li Hongmei, Zhang Chuansong, Han Xiurong, et al. Changes in concentrations of oxygen, dissolved nitrogen, phosphate, and silicate in the southern Yellow Sea, 1980−2012: Sources and seaward gradients[J]. Estuarine, Coastal and Shelf Science, 2015, 163: 44−55. -
下载:
图(4) / 表(1)
计量
- 文章访问数: 571
- HTML全文浏览量: 78
- PDF下载量: 150
- 被引次数: 0
