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49 5 Vol.49, No.5 2018 9 OCEANOLOGIA ET LIMNOLOGIA SINICA Sep., 2018 : * 1, 2, 3, 4 1, 2, 3, 4 1, 2, 4 1 (1. ( ) 266071; 2. 266071; 3. 100049; 4. 266071) 2007, (green tides), 2018 12,,,,, (Ulva prolifera),,,,,,,, 黄海 ; 绿潮 ; 成因 ; 防控 Q14 doi: 10.11693/hyhz20180700158 (green tide),,,,,,, (Smetacek et al, 2013; Gao et al, 2017) 2007 6,, 6000 2008 6,,,, 60 1/3 (, 2008),,, 80,, 12 ( 1),,,,,, *, 41676106 ;, 2016ASKJ02 ; (A ), XDA11020304 ; -, U1606404,,, E-mail: rcyu@qdio.ac.cn : 2018-07-04, : 2018-08-22

5 : : 943, (Sun et al, 2008;, 2010; Keesing et al, 2011; Liu et al, 2013a, b; Wang et al, 2015; Zhou et al, 2015;, 2018),,, 1 2007 Fig.1 Variation of sea area affected or covered by green tides in the Yellow Sea from 2007 ( ) 1 2008,,,,, (Ulva prolifera);,, ; (Sun et al, 2008),, (, 2008),,,, (Liu et al, 2009; Pang et al, 2010; Zhang et al, 2011; Liu et al, 2013b) 2009, 973,,, (Zhou et al, 2015; Wang et al, 2015),, (Huo et al, 2016;, 2017),, 1.1,,, (, 2008;, 2009; Ding et al, 2009; Leliaert et al, 2009),,

944 49,, RNA (rdna) (ITS),, ITS (Ulva linza U. procera), ITS 5S rdna (Shimada et al, 2008; Duan et al, 2012),, DNA (ITS 5SrDNA ), (Zhao et al, 2013);, (Zhao et al, 2011) DNA,,, (Xiao et al, 2013; Zhang et al, 2015a),,, (Zhang et al, 2015b), ( ) DNA,, (Zhao et al, 2011, 2015), DNA PCR (Zhao et al, 2015) 1.2,, (, 2008; Liu et al, 2009; Hu et al, 2010; Xu et al, 2014; Qi et al, 2016),, (Liu et al, 2009, 2013b; Pang et al, 2010; Zhang et al, 2011), (Keesing et al, 2011; Zhang et al, 2015b; Huo et al, 2016),,,, (Liu et al, 2009; Hu et al, 2010), (Liu et al, 2010; Zhang et al, 2015a), (Zhang et al, 2018),,,,, 5 (Liu et al, 2010; Li et al, 2015a; Zhang et al, 2015b),,, ( 2),, (Liu et al, 2012), (Liu et al, 2012; Huo et al, 2014; Li et al, 2014),, 15 o C (Song et al, 2015),, 5, (Li et al, 2015a) 4 5,,,, (Gao et al, 2016),,,,,,, ( 3),

5 : : 945 2 ( Zhou et al, 2015) Fig.2 Illustration of the early development stage of green tide in Subei Shoal (modified from Zhou et al, 2015) Fig.3 3 Illustration of the floating routes of green algae in Subei Shoal 5,,,, 1.3,,,,,, (Liu et al, 2015a),,,,, (Li et al, 2016a, b),, (Xu et al, 2012),, 10% 37%, (Gao et al, 2016),,,,,,,,, (, 2012),,, (Liu et al, 2017; Li et al, 2017), (Liu et al, 2015b; Shi et al, 2015),,, (Bao et al, 2015),,,,,

946 49, (Liu et al, 2015b),, (Geng et al, 2015;, 2018), 20 000, 6 10 7 m (Geng et al, 2015),, (Liu et al, 2010; Wang et al, 2015;, 2017),, 2005 2009,,, 2010 (, 2017) 1.4,,,,,, 2008 20 (Ye et al, 2011),,, (Wang et al, 2011) 2008, 8, 3 4,, (Xing et al, 2015),,, (Wang et al, 2012;, 2018) 2 2007,,,,,,,, (Zhao et al, 2015),,,,,,,,,,,, (Xu et al, 2014; Qi et al, 2016),,, 2007,,,, 1999, 2006,,

5 : : 947,, N/P (Li et al, 2015b),,,,,,,,,,,,,, 2016,,,,, ;,,,,, (, 2018) 3 2007,,,,,,,,,,,, 2009. (Enteromorpha prolifera)., 40(1): 68 71,,, 2018.., 40(2): 1 13, 2008.,., (11): 37 40,,, 2012.., 34(4): 147 154,,, 2018. 2017 35 N., 49(5): 1021 1030,,, 2008. 2008., 26(3): 409 410,,, 2017.., 41(2): 35 43,,, 2008.., 15(5): 822 829,,, 2018.., 49(5): 1006 1013,,, 2010.., (1): 5 9,,, 2018.., 49(5): 950 958 Bao M, Guan W B, Yang Y et al, 2015. Drifting trajectories of green algae in the western Yellow Sea during the spring and summer of 2012. Estuarine, Coastal and Shelf Science, 163: 9 16 Ding L P, Fei X G, Lu Q Q et al, 2009. The possibility analysis of habitats, origin and reappearance of bloom green alga (Enteromorpha prolifera) on inshore of western Yellow Sea. Chinese Journal of Oceanology and Limnology, 27(3): 421 424 Duan W J, Guo L X, Sun D et al, 2012. Morphological and molecular characterization of free-floating and attached green macroalgae Ulva spp. in the Yellow Sea of China. Journal of Applied Phycology, 24(1): 97 108 Gao G, Clare A S, Rose C, 2017. Eutrophication and warming-driven green tides (Ulva rigida) are predicted to increase under future climate change scenarios. Marine Pollution Bulletin, 114(1): 439 447 Gao G, Zhong Z H, Zhou X H et al, 2016. Changes in morphological plasticity of Ulva prolifera under different environmental conditions: a laboratory experiment. Harmful Algae, 59: 51 58 Geng H X, Yan T, Zhou M J et al, 2015. Comparative study of the germination of Ulva prolifera gametes on various substrates. Estuarine, Coastal and Shelf Science, 163: 89 95 Hu C M, Li D Q, Chen C S et al, 2010. On the recurrent Ulva prolifera blooms in the Yellow Sea and East China Sea. Journal of Geophysical Research: Oceans, 115(C5): C05017

948 49 Huo Y Z, Han H B, Hua L et al, 2016. 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. Harmful Algae, 59: 91 99 Huo Y Z, Hua L, Wu H L et al, 2014. Abundance and distribution of Ulva microscopic propagules associated with a green tide in the southern coast of the Yellow Sea. Harmful Algae, 39: 357 364 Keesing J K, Liu D Y, Fearns P et al, 2011. Inter- and intra-annual patterns of Ulva prolifera green tides in the Yellow Sea during 2007 2009, their origin and relationship to the expansion of coastal seaweed aquaculture in China. Marine Pollution Bulletin, 62(6): 1169 1182 Leliaert R, Zhang X W, Ye N H et al, 2009. Research note: identity of the Qingdao algal bloom. Phycological Research, 57(2): 147 151 Li H M, Zhang C S, Han X R et al, 2015b. Changes in concentrations of oxygen, dissolved nitrogen, phosphate, and silicate in the southern Yellow Sea, 1980 2012: sources and seaward gradients. Estuarine, Coastal and Shelf Science, 163: 44 55 Li H M, Zhang Y Y, Han X R et al, 2016b. Growth responses of Ulva prolifera to inorganic and organic nutrients: implications for macroalgal blooms in the southern Yellow Sea, China. Scientific Reports, 6: 26498 Li H M, Zhang Y Y, Tang H J et al, 2017. Spatiotemporal variations of inorganic nutrients along the Jiangsu coast, China, and the occurrence of macroalgal blooms (green tides) in the southern Yellow Sea. Harmful Algae, 63: 164 172 Li S X, Yu K F, Huo Y Z et al, 2016a. Effects of nitrogen and phosphorus enrichment on growth and photosynthetic assimilation of carbon in a green tide-forming species (Ulva prolifera) in the Yellow Sea. Hydrobiologia, 776(1): 161 171 Li Y, Song W, Xiao J et al, 2014. Tempo-spatial distribution and species diversity of green algae micro-propagules in the Yellow Sea during the large-scale green tide development. Harmful Algae, 39: 40 47 Li Y, Xiao J, Ding L P et al, 2015a. Community structure and controlled factor of attached green algae on the Porphyra yezoensis aquaculture rafts in the Subei Shoal, China. Acta Oceanologica Sinica, 34(8): 93 99 Liu D Y, Keesing J K, Dong Z J et al, 2010. Recurrence of the world s largest green-tide in 2009 in Yellow Sea, China: Porphyra yezoensis aquaculture rafts confirmed as nursery for macroalgal blooms. Marine Pollution Bulletin, 60(9): 1423 1432 Liu D Y, Keesing J K, He P M et al, 2013a. The world's largest macroalgal bloom in the Yellow Sea, China: Formation and implications. Estuarine, Coastal and Shelf Science, 129: 2 10 Liu D Y, Keesing J K, Xing Q G et al, 2009. World s largest macroalgal bloom caused by expansion of seaweed aquaculture in China. Marine Pollution Bulletin, 58(6): 888 895 Liu F, Pang S J, Chopin T et al, 2013b. Understanding the recurrent large-scale green tide in the Yellow Sea: Temporal and spatial correlations between multiple geographical, aquacultural and biological factors. Marine Environmental Research, 83: 38 47 Liu F, Pang S J, Zhao X B et al, 2012. Quantitative, molecular and growth analyses of Ulva microscopic propagules in the coastal sediment of Jiangsu province where green tides initially occurred. Marine Environmental Research, 74: 56 63 Liu J A, Su N, Wang X L et al, 2017. Submarine groundwater discharge and associated nutrient fluxes into the Southern Yellow Sea: A case study for semi enclosed and oligotrophic seas implication for green tide bloom. Journal of Geophysical Research: Oceans, 122(1): 139 152 Liu Q, Yu R C, Yan T et al, 2015a. Laboratory study on the life history of bloom-forming Ulva prolifera in the Yellow Sea. Estuarine, Coastal and Shelf Science, 163: 82 88 Liu X Q, Li Y, Wang Z L et al, 2015b. Cruise observation of Ulva prolifera bloom in the southern Yellow Sea, China. Estuarine, Coastal and Shelf Science, 163: 17 22 Pang S J, Liu F, Shan T F et al, 2010. Tracking the algal origin of the Ulva bloom in the Yellow Sea by a combination of molecular, morphological and physiological analyses. Marine Environmental Research, 69(4): 207 215 Qi L, Hu C M, Xing Q G et al, 2016. Long-term trend of Ulva prolifera blooms in the western Yellow Sea. Harmful Algae, 58: 35 44 Shi X Y, Qi M Y, Tang H J et al, 2015. Spatial and temporal nutrient variations in the Yellow Sea and their effects on Ulva prolifera blooms. Estuarine, Coastal and Shelf Science, 163: 36 43 Shimada S, Yokoyama N, Arai S et al, 2008. Phylogeography of the genus Ulva (Ulvophyceae, Chlorophyta), with special reference to the Japanese freshwater and brackish taxa. Journal of Applied Phycology, 20(5): 979 989 Smetacek V, Zingone A, 2013. Green and golden seaweed tides on the rise. Nature, 504(7478): 84 88 Song W, Peng K Q, Xiao J et al, 2015. Effects of temperature on the germination of green algae micro-propagules in coastal waters of the Subei Shoal, China. Estuarine, Coastal and Shelf Science, 163: 63 68 Sun S, Wang F, Li C L et al, 2008. Emerging challenges: massive green algae blooms in the Yellow Sea. Nature Proceedings, NPRE20082266-1 Wang C, Yu R C, Zhou M J, 2011. Acute toxicity of live and decomposing green alga Ulva (Enteromorpha) prolifera to abalone Haliotis discus hannai. Chinese Journal of Oceanology and Limnology, 29(3): 541 546 Wang C, Yu R C, Zhou M J, 2012. Effects of the decomposing green macroalga Ulva (Enteromorpha) prolifera on the growth of four red-tide species. Harmful Algae, 16: 12 19 Wang Z L, Xiao J, Fan S L et al, 2015. 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. Limnology and Oceanography, 60(4): 1105 1117

5 : : 949 Xiao J, Li Y, Song W et al, 2013. Discrimination of the common macroalgae (Ulva and Blidingia) in coastal waters of Yellow Sea, northern China, based on restriction fragment-length polymorphism (RFLP) analysis. Harmful Algae, 27: 130 137 Xing Q G, Hu C M, Tang D L et al, 2015. World's largest macroalgal blooms altered phytoplankton biomass in summer in the Yellow Sea: satellite observations. Remote Sensing, 7(9): 12297 12313 Xu J F, Fan X, Zhang X W et al, 2012. Evidence of coexistence of C 3 and C 4 photosynthetic pathways in a green-tideforming alga, Ulva prolifera. PLoS One, 7(5): e37438 Xu Q, Zhang H Y, Ju L et al, 2014. Interannual variability of Ulva prolifera blooms in the Yellow Sea. International Journal of Remote, 35(11 12): 4099 4113 Ye N H, Zhang X W, Mao Y Z et al, 2011. Green tides are overwhelming the coastline of our blue planet: taking the world's largest example. Ecological Research, 26(3): 477 Zhang Q C, Liu Q, Kang Z J et al, 2015a. Development of a fluorescence in situ hybridization (FISH) method for rapid detection of Ulva prolifera. Estuarine, Coastal and Shelf Science, 163: 103 111 Zhang Q C, Liu Q, Yu R C et al, 2015b. Application of a fluorescence in situ hybridization (FISH) method to study green tides in the Yellow Sea. Estuarine, Coastal and Shelf Science, 163: 112 119 Zhang Q C, Yu R C, Chen Z F et al, 2018. Genetic evidence in tracking the origin of Ulva prolifera blooms in the Yellow Sea, China. Harmful Algae, 78: 86 94. Zhang X W, Xu D, Mao Y Z et al, 2011. Settlement of vegetative fragments of Ulva prolifera confirmed as an important seed source for succession of a large-scale green tide bloom. Limnology and Oceanography, 56(1): 233 242 Zhao J, Jiang P, Liu Z Y et al, 2011. Genetic variation of Ulva (Enteromorpha) prolifera (Ulvales, Chlorophyta) the causative species of the green tides in the Yellow Sea, China. Journal of Applied Phycology, 23(2): 227 233 Zhao J, Jiang P, Liu Z Y et al, 2013. The Yellow Sea green tides were dominated by one species, Ulva (Enteromorpha) prolifera, from 2007 to 2011. Chinese Science Bulletin, 58(19): 2298 2302 Zhao J, Jiang P, Qin S et al, 2015. Genetic analyses of floating Ulva prolifera in the Yellow Sea suggest a unique ecotype. Estuarine, Coastal and Shelf Science, 163: 96 102 Zhou M J, Liu D Y, Anderson D M et al, 2015. Introduction to the special issue on green tides in the Yellow Sea. Estuarine, Coastal and Shelf Science, 163: 3 8 PROGRESSES AND PERSPECTIVES ON GREEN-TIDE STUDIES IN THE YELLOW SEA YU Ren-Cheng 1, 2, 3, 4, SUN Song 1, 2, 3, 4, YAN Tian 1, 2, 4, ZHOU Ming-Jiang 1 (1. CAS Key Laboratory of Marine Ecology and Environmental Sciences, Institute of Oceanology, Chinese Academy of Sciences, Qingdao 266071, China; 2. Laboratory for Marine Ecology and Environmental Science, Qingdao National Laboratory for Marine Science and Technology, Qingdao 266071, China; 3. University of Chinese Academy of Sciences, Beijing 100049, China; 4. Center for Ocean Mega-Science, Chinese Academy of Sciences, Qingdao 266071, China) Abstract Large-scale green tides, which have been recorded in the Yellow Sea for 12 years since 2007, posed significant impacts on the scenery, environment and mariculture industry along the west coast of the southern Yellow Sea. Green tides have been considered as a routine marine ecological disaster in the Yellow Sea, and regional governments of Jiangsu and Shandong provinces have to put huge manpower and resources every year to collect and manage the green algae accumulated along the coastline. Targeting on the green tides in the Yellow Sea, Chinese government organized scientists to work together intensively on the origin, mechanisms, impacts, monitoring and control of green tides during the last decade, and significant progresses have been made on the bloom-forming species, origin of green tides, and key factors affecting the green tides. The major bloom-forming green alga has been identified as Ulva prolifera, and the origin of green tides have been traced back to the Subei Shoal along the west coast of the southern Yellow Sea. Major processes related to the early development of green tides in Subei Shoal have been elucidated. The biological characteristics, unique environmental features of Subei Shoal, as well as the intensive mariculture activities in Subei Shoal are key factors leading to the formation of green tides. Some questions, however, still need to be answered in future studies, such as the origin of unique bloom-forming U. prolifera, the long-term trend of green tides in the Yellow Sea, the ecological consequences of recurrent green tides, and the prevention and control strategies against green tides etc. Focusing on the green tide issue in the Yellow Sea, a project "mechanisms and prevention strategies on marine ecological disasters " was supported recently by the Qingdao National Laboratory for Marine Science and Technology to test the ideas on the formation of green tides, and to offer knowledge and techniques on the monitoring and prevention strategies on green tides. Key words Yellow Sea; green tide; mechanism; prevention