한수지 49(5), 657-664, 2016 Original Artile Korean J Fish Aquat Si 49(5),657-664,2016 녹조대발생종금발대마디말 (Cladophora vadorum) 의절편생장에온도, 조도및영양염종류가미치는영향 나연주 전다빈 이정록 김영식 1 최한길 * 남기완 2 원광대학교생명과학부 / 환경과학연구소, 1 군산대학교해양생물공학과, 2 부경대학교자원생물학과 Effets of Temperature, Irradiane, and Nutrient Type on the Fragment Growth of Green Tide Alga Cladophora vadorum Yeon Ju Na, Da Vine Jeon, Jung Rok Lee, Young Sik Kim 1, Han Gil Choi* and Ki Wan Nam 2 Faulty of Biologial Siene and Institute for Environmental Siene, Wonkwang University, Iksan 54538, Korea 1 Department of Marine Biotehnology, Kunsan National University, Kunsan 54150, Korea 2 Department of Marine Biology, Pukyong National University, Busan 48513, Korea The green maroalga Cladophora vadorum loomed along the oast at Sangrok Beah, Buan, South Korea, in Septemer 2015. To eluidate the ause of loom, the effets of environmental fators on the vegetative growth of adult fragments were examined. Growth experiments were arried out under different ominations of temperatures and irradianes, and with a single fator of nutrients (nitrogen, phosphorus). The maximal growth of C. vadorum was reported under the omination of 25 C and 100 μmol photons m -2 s -1. The speies grew under a wide range of N and P onentrations. The growth of C. vadorum peaked at 50 μm PO 4 3-, 80 μm NH 4+, and 100 μm NO 3-. Adult fragments formed holdfasts and new ranhes within 3 days in ulture and eame adults, showing polarized growth patterns, in 2 weeks. This is the first report showing the development of numerous ladelets from a segment in Cladophora speies. The present results indiate that Cladophora looms appear under growth onditions that are favorale in terms of temperatures, irradiane, and nutrients via fragment growth patterns produing rapid holdfasts and many ladelets. Key words: Cladophora vadorum, Fragmentation, Eutrophiation, Temperature, Green tide 서론 (green tides) (Charlier et al., 2007; Shimada et al., 2008; Zhang et al., 2013), (Blomster et al., 2002; Largo et al., 2004; Ye et al., 2011). (Ulva), (Chaetomorpha), (Cladophora), (Rhizolonium) (Flether, 1996; Taylor et al., 2001; Nelson et al., 2003), (Nelson and Lee, 2001; Franz and Friedman, 2002)., (seagrass ed) (Råerg et al., 2005), ( ), (dead zone) (Hallegraeff, 1993; Worm et al., 2001).,,, (Liu et al., 2009; Hu et al., 2010; Ye et al., 2011). ( ) 2008 (Ulva prolifera) 100, 1,000 (US$ 100 million) (Wang et al., 2009), http://dx.doi.org/10.5657/kfas.2016.0657 Korean J Fish Aquat Si 49(5) 657-664, Otoer 2016 This is an Open Aess artile distriuted under the terms of the Creative Commons Attriution Non-Commerial Liens (http://reativeommons.org/lienses/y-n/3.0/) whih permits unrestrited non-ommerial use, distriution, and reprodution in any medium, provided the original work is properly ited. Reeived 18 July 2016; Revised 19 Septemer 2016; Aepted 27 Septemer 2016 *Corresponding author: Tel: +82. 63. 850. 6579 Fax: +82. 63. 857. 8837 E-mail address: hghoi@wku.a.kr Copyright 2016 The Korean Soiety of Fisheries and Aquati Siene 657 pissn:0374-8111, eissn:2287-8815
658 나연주ㆍ전다빈ㆍ이정록ㆍ김영식ㆍ최한길ㆍ남기완 (Zhang et al., 2014; Wang et al., 2015).,, (Zhang et al., 2013),., (Pekol et al., 1994; Duarte, 1995; Tan et al., 1999; Mereron et al., 2007; Nelson et al., 2008; Choi et al., 2010), (zoosporangia) (gametangia), (Dan et al., 1997; Deng et al., 2011; Zhang et al., 2013, 2016)., Ulva mutailis U. prolifera (= Enteromorpha prolifera) (Nordy and Hoxmark, 1972; Dan et al., 1997; Lin et al., 2008), Ulva prolifera (fragment) (Zhang et al., 2016). (fragment regeneration) (Bryoposis hypnoide) Chaetomorpha valida, (Deng et al., 2011). (Cladophora spp.) 214,, (Dodds and Gudder, 1992; Graham et al., 2009; Guiry and Guiry, 2016).,, (ph) (Entwisle, 1989; Taylor et al., 2001), (,, ) (Pitkänen et al., 1997; Vahteri et al., 2000; Higgins et al., 2008). Cladophora vagaunda (N, nitrogen) (P, phosphorus), (Pekol et al., 1994)., (Higgins et al., 2008) (Valiela et al., 1997; Lomstein et al., 2006; Balkis et al., 2013). (Cladophora glomerata), (Higgins et al., 2008)., 2015 9 214 (Cladophora vadorum)., Ha et al. (2016) (0.438 mg L -1 ) (0.046 mg L -1 ).,,. 재료및방법 (34 60' N, 126 48' E) 2015 9,, (0.45 m)., (3 g) (100 ml) 8,000 rpm 10 s (Ø 0.5 mm) 3-4 (fragments). 성체의형태와발생, 6 (2.5 2.5 m) 15 ml PES (Provasoli, 1968) (Ø 9 m) 3 ml (1 ml 100 ), 25, 100 µmol photons m -2 s -1 12:12h L:D (Light:Dark) (Bionex, Korea) 2. 3, (DX- 200). (Olympus BH-2, Japan) (Canon IXUS 860IS). 온도와조도영향, PES (5 ml) (1 ml) (15, 20, 25, 30 ) (50, 100 mol photons m -2 s -1 ) 8. 12:12h L:D,. 3 GeO 2 (5 mgl -1 ) (Shea and Chopin, 2007). 8,. 3, 30 Image J (RGR,
녹조대발생종금발대마디말성체의절편생장 659 relative growth rate) (Serisawa et al., 2002). RGR (% day -1 ) = 100 (In L 2 In L 1 )/T 2 T 1 L 2, L 1 T 2 T 1 (day). 영양염종류와생장, (100 ml) 3 NaH 2 PO 4 (0.142 g) stok solution (10 mm), NH 4 Cl (0. 540 g) stok solution (100 mm), NaNO 3 (0.850 g, Sigma Co., USA) stok solution (100 mm)., (stok solution) (0, 10, 20, 50, 100 M), (0, 5, 10, 40, 80, 100 M) (0, 10, 40, 80, 100, 200 M). (low nutrient seawater, OSIL, LNS23, UK) (nitrate), (nitrite), (phosphate) <0.1 M (siliate) <0.3 M., (5 ml) (1 ml) 3. 3, 25, 100 mol photons m -2 s -1 12:12h L:D 6.. 통계분석 (,, ), Cohran s test (homogeneity of varianes), one-way ANOVA test ( ) two way ANOVA test ( ). Tukey s HSD test (Sokal and Rohlf, 1995), STATISTICA version 7.0. 성체의형태와발생 결과, 183.94 5.96 m (mean SE, n=50 ells), 31.28 1.09 m (n=50) / 6.01 0.17 m (Fig. 1A, 1B)., (23.74 0.67 m, n=50) (Fig. 1C). 3,, 2 (Fig. 1D, 1E). 4 0.02 0.00 mm 2 (Fig. 1F)., (Fig. 1C), (Fig. 1G). 8 (Fig. 1H, 1I). 온도와조도영향 - 320.79 8.71 m (mean SE, n=300 fragments), 8 556.17-2358.71 m. 1,442.08 178.69 m (50 mol photons m -2 s -1 ) 1,624.46 149.67 m (100 mol photons m -2 s -1 ) (F 1,16 =46.72, P<0.01). (15-30 ), 15 762.75 93.50 m 25 2,288.81 40.31 m, 25 (P<0.01), 20 30 (Tukey test, P 0.01, Tale 1). 8.62-32.53 % day -1 25 100 mol photons m -2 s -1 (Fig. 2). 영양염종류와생장, 377.27 9.58 m Tale 1. Analysis of variane (two-way ANOVA) for the effets of temperature and irradiane on relative growth rate (RGR, % day -1 ) of Cladophora vadorum after 8 days in ulture Fators df MS F P Temperature (T) 3 2330732.5 546.4 < 0.001 Irradiane (I) 1 199290.4 46.7 < 0.001 Interation 3 37762.5 8.9 < 0.01 Error 16 4265.4 Tukey test (P =0.05) Temperature 15<20=30<25 Irradiane 50<100 df, dgrees of freedom; MS, mean square; F, F-statisti variane ratio; P, proaility.
660 나연주ㆍ전다빈ㆍ이정록ㆍ김영식ㆍ최한길ㆍ남기완 Fig. 1. Development of new filaments of Cladophora vadorum. A: Field-olleted materials. B: Cut fragments. C: A developed new filaments (arrow). D: Formation of holdfast. E: A developed holdfast after 2 weeks in ulture. F: Holdfast development after 4 weeks. G: Two parts of holdfasts. H: Seondary ranhes from holdfast after 8 weeks of inuation. I: 8 weeks old ultured fragments. Sale ars are 200 µm (A,G), 100 µm (B-C), 50 μm (D-F), 1 mm (H), 0.2 m (I). RGR (% day -1 ) 40 30 20 10 0 50 100 μmol photons m -2 s -1 15 20 25 30 Temperature ( ) Fig. 2. Relative growth rate (RGR, % day -1 ) of Cladophora vadorum ultured under various temperature and irradiane levels for 8 days. Culture onditions were 34 psu and 12:12h L:D. Data represents mean±se (n=3 repliates) (mean SE, n=300), 6, (0, 10, 20, 50, 100 M) 839.32-978.74 m. 12.74-15.42% day -1 50 M (100 M) (Fig. 3A)., 3 10, 20, 100 M (Tukey HSD test, P 0.01). 5-100 M, 6 826.93-1,019.30 m 12.69-16.08% day -1 (Fig. 3B). 80 M 100 M (F 5,12 =104.49, P<0.01). 6 751.97-1035.34 m 11.26-16.47% day -1 (Fig. 3C). (0, 5, 10, 40 M) (80, 100 M) (F 6,14 =99.13, P<0.01), 200 M. 80 100 M,, (0, 5 M) 10, 40 200 M (Tukey test, P 0.01).
녹조대발생종금발대마디말성체의절편생장 661 (A) (B) (C) RGR (% day -1 ) RGR (% day -1 ) RGR (% day -1 ) 20 16 12 8 4 0 20 16 12 8 4 0 20 16 12 8 4 0 a a a 0 10 20 50 100 PO 4 (μmol/l) 0 5 10 40 80 100 a NH 4 (μmol/l) 0 5 10 40 80 100 200 NO 3 (μmol/l) Fig. 3. Relative growth rate (RGR, % day -1 ) of Cladophora vadorum grown in PO 4 3- (A), NH 4 + (B) and NO 3 - (C) for 6 days. Fragments were ultured under 25, 100 μmol photons m -2 s -1 and 12:12h L:D. Vertial ars indiate standard errors (n=3 repliates). Different letters indiate signifiant group of mean found with the Tukey HSD test. 고찰 d,, (Taylor et al., 2001; Nelson et al., 2008). 15 20 25, 2015 8-9 25., (Cladophora alida) (C. vagaunda) 25 Breeman et al. (2002). (50 mol photons m -2 s -1 ) 100 mol photons m -2 s -1, (20 120 mol photons m -2 s -1 )., 20-30 25-33% day -1 10-25% day -1 (16-26, 100-200 mol photons m -2 s -1 ) (Luo et al., 2012),., (regeneration) (Maggs and Stegenga, 1999; Boudouresque and Verlaque, 2002; Nyerg and Wallentinus, 2005)., (vegetative propagation) (Lin et al., 2008), (fragment regeneration) (Hiraoka et al., 2004; Deng et al., 2011). 1992-2000 ( ) 2008 ( ) (Blomster et al., 2002)., (Ulva intestinalis, U. ompressa) (lade-let) (Zhang et al., 2016). (polarized growth). (Eaton et al., 1966; Lee and Wihroski, 1996; Zhang et al., 2016),. (0 M) (10-100 M), (5-100 M), (5-200 M), 50 M ( ), 80 M ( ) 100 M ( ). (Cladophora dalmatia) 10 M ( ), 100 M ( ) 800 M ( ) 4-8% day -1 (Taylor et al., 2001), Cladophora oelothrix 35-70 M 10-15% day -1 (de Paula Silva et al., 2008).
662 나연주ㆍ전다빈ㆍ이정록ㆍ김영식ㆍ최한길ㆍ남기완 15-17% day -1 (4-8% day -1 ) Cladophora oelothrix (10-15% day -1 ). 1970 1990 (210-421 M) (Niholls et al., 2001), ( ) (Higgins et al., 2005a, 2005)., Taylor et al. (2001) 30 M 100 M, (100 M), (100 M), (200 M). (Sousa et al., 2007; Liu et al., 2010),, (Kim et al., 2004; Liu et al., 2009; Zhao et al., 2011). 2015 8-9, Ha et al. (2016) (TP) (5 M) (TN) (30 M) 10 ( ) 5 ( ),., 2015 Ha et al. (2016), (,, )., (lade-let).,.,. 사사.,. Referenes Balkis N, Sivri N, Linda F, Muharrem B, Turgay D and Atakan S. 2013. Exessive growth of Cladophora (Dillwyn) kützing and enteri ateria in mats in the Southwestern Istanul oast, Sea of Marmama. IUFS J Biol 72, 43-50. Blomster J, Bäk S, Fewer DP, Kiirikki M, Lehvo A, Maggs CA and Stanhope MJ. 2002. Novel morphology in Enteromorpha (Ulvophyeae) forming green tides. Am J Bot 89, 1756-1763. http://dx.doi.org/10.3732/aj.89.11.1756. Boudouresque CF and Verlaque M. 2002. Biologial pollution in the Mediterranean Sea: invasive versus introdued marophytes. Mar Pollut Bull 44, 32-38. http://dx.doi.org/10.1016/ S0025-326X(01)00150-3. Breeman A, Oh YS, Hwang MS and Van Den Hoek C. 2002. Evolution of temperature responses in the Cladophora vagaunda omplex and the C. alida/seriea omplex (Chlorophyta). Eur J Phyol 37, 45-58. http://dx.doi.org/10. 1017/S0967026201003420. Charlier RH, Morand P, Finkl CW and Thys A. 2007. Green tides on the Brittany oasts. Environ Res Eng Manage 3, 52-59. http://dx.doi.org/10.1109/baltic.2006.7266128. Choi TS, Kang EJ, Kim JH and Kim KY. 2010. Effets of salinity on growth and nutrient uptake of Ulva pertusa (Chlorophyta) from an eelgrass ed. Algae 25, 17 26. http:// dx.doi.org/10.4490/algae.2010.25.1.017. Dan A, Ohno M and Matuoka M. 1997. Cultivation of the green alga Enteromorpha prolifera using hopped tissue for artifiial seeding. Suisan Zoshoku 45, 5-8. de Paula Silva PH, MBride S, de Nys R and Oaul NA. 2008. Integrating filamentous green tide algae into tropial pondased aquaulture. Aquaulture 284, 74-80. http://dx.doi. org/10.1016/j.aquaulture.2008.07.035. Deng YY, Tang XR, Huang BX and Ding LP. 2011. Life history of Chaetomorpha valida (Cladophoraeae, Chlorophyta) in ulture. Bot Mar 54, 551-556. http://dx.doi.org/10.1515/ BOT.2011.066. Dodds WK and Gudder DA. 1992. The eology of Cladophora. J Phyol 28, 415-427. http://dx.doi.org/10.1111/j.0022-3646.1992.00415.x. Duarte C. 1995. Sumerged aquati vegetation in relation to different nutrient regimes. Ophelia 41, 87-112. http://dx.doi.or g/10.1080/00785236.1995.10422039. Eaton JW, Brown JG and Rouno FE. 1966. Some oservations on polarity and regeneration in Enteromorpha. Br Phyol Bull 3, 53-62. http://dx.doi.org/10.1080/00071616600650071. Entwisle TJ. 1989. Phenology of the Cladophora-Stigeolonium ommunity in two uran reeks of Melourne. Aust J Mar Freshwater Res 40, 471-489. http://dx.doi.org/10.1071/ MF9890471. Flether RL. 1996. The British Isles. In: Sharmm W, Nienhuis PH(eds) Marine enthi vegetion: reent hanges, the ef-
녹조대발생종금발대마디말성체의절편생장 663 fets of eutrophiation. Springer, Berlin, pp 150-223. Franz DR and Friedman I. 2002. Effets of a maroalgal mat (Ulva latua) on estuarine sand flat opepods: and experimental study. J Exp Mar Biol Eol 271, 209-226. http:// dx.doi.org/10.1016/s0022-0981(02)00045-x. Graham LE, Graham JM and Wilox LW. 2009. Algae, 2nd ed. Pearson, San Franiso. Guiry MD and Guiry GM. 2016. AlgaeBase. World-wide eletroni puliation, National University of Ireland, Galway. http://www.algaease.org. Ha DS, Yoo HI, Chang SJ and Hwang EK. 2016. Bloom of a filamentous green alga Cldophora vadorum (Areshoug) Kützing and nutrient levels at Shangrok eah, Buan, Korea. Korean J Fish Aquat Si 49, 241-246. http://dx.doi. org/10.5657/kfas.2016.0241. Hallegraeff GM. 1993. A review of harmful algal looms and their apparent gloal inrease. Phyologia 32, 79-99. http:// dx.doi.org/10.2216/i0031-8884-32-2-79.1. Higgins SN, Heky RE and Guildford SJ. 2005a. Modeling the growth, iomass, and tissue phosphorus onentration of Cladophora glomerata in eastern Lake Erie: model desription and field testing. J Great Lakes Res 31, 439-455. http:// dx.doi.org/10.1016/s0380-1330(05)70275-6. Higgins SN, Howell TE, Hekky RE, Guildford SJ and Smith RE. 2005. The wall of green: the status of Cladophora on the northern shores of Lake Erie s eastern asin, 1995-2002. J Great Lakes Res 31, 547-563. http://dx.doi.org/10.1016/ S0380-1330(05)70283-5. Higgins SN, Malkin SY, Howell ET, Guildford SJ, Campell L, Hiriart-Baer V and Heky RE. 2008. An eologial review of Cladophora glomerata (Chlorophyta) in the Laurentian Great Lakes. J Phyol 44, 839-854. http://dx.doi. org/10.1111/j.1529-8817.2008.00538.x. Hiraoka M, Ohno M, Kawaguhi S and Yoshida G. 2004. Crossing test among floating Ulva thalli forming greentide in Japan. Hydroiologia 512, 239-245. http://dx.doi. org/10.1023/b:hydr.0000020332.12641.a2. Hu C, Li D, Chen C, Ge J, Muller-Karger FE, Liu J, Yu F and He MX. 2010. On the reurrent Ulva prolifera looms in the Yellow Sea and East China Sea. J Geophys Res 79, 185-194. http://dx.doi.org/10.1029/2009jc005561. Kim KY, Choi TS, Kim JH, Han T, Shin HW and Garary DJ. 2004. Physiologial eology and seasonality of Ulva pertusa on a temperate roky shore. Phyologia 43, 483-492. http://dx.doi.org/10.2216/i0031-8884-43-4-483.1. Largo DB, Semrano J, Hiraoka M and Ohno M. 2004. Taxonomi and eologial profile of green tide speies of Ulva (Ulvales, Chlorophyta) in entral Philippines. Hydroiologia 512, 247-253. http://dx.doi.org/10.1007/978-94-007-0944-7_32. Lee TF and Wihroski M. 1996. Polar regeneration in Enteromorpha prolifera. J Phyol 32, 27. Lin A, Shen S, Wang J and Yan B. 2008. Reprodution diversity of Enteromorpha prolifera. J Integr Plant Biol 50, 622-629. http://dx.doi.org/10.1111/j.1744-7909.2008.00647.x. Liu D, Keesing JK, Xing Q and Shi P. 2009. World s largest maroalgal loom aused y expansion of seaweed aquaulture in China. Mar Pollut Bull 58, 888-895. http://dx.doi. org/10.1016/j.marpolul.2009.01.013. Liu D, Keesing JK, Dong Z, Zhen Y, Di B, Shi Y, Fearns P and Shi P. 2010. Reurrene of the world s largest green-tide in 2009 in Yellow Sea, China: Porphyra yezoensis aquaulture rafts onfirmed as nursery for maroalgal looms. Mar Pollut Bull 60, 1423-1432. http://dx.doi.org/10.1016/j.marpolul.2010.05.015. Lomstein BA, Gulderg LB, Neuauer AA, Hansen J, Donnelly A, Herert RA, Viaroli P, Giordani G, Azzoni R, Wit R and Finster K. 2006. Benthi deomposition of Ulva latusa: a ontrolled laoratory experiment. Aquat Bot 85, 271-281. http://dx.doi.org/10.1016/j.aquaot.2006.05.006. Luo MB, Liu F and Xu ZL. 2012. Growth and nutrient uptake apaity of two o-ourring speies, Ulva prolifera and Ulva linza. Aquat Bot 100, 18-24. http://dx.doi.org/10.1016/j. aquaot.2012.03.006. Maggs CA and Stegenga H. 1999. Red algal exoti on North Sea oasts. Helgoländer Meeresun 52, 243-258. http://dx.doi. org/10.1007/bf02908900. Mereron M, Antoine V, Auy I and Morand P. 2007. In situ growth potential of the sutidal part of green tide forming Ulva spp. stoks. Si Total Environ 384, 293-305. http:// dx.doi.org/10.1016/j.sitotenv.2007.05.007. Nelson TA and Lee A. 2001. A manipulative experiment demonstrates that looms of the maroalga Ulvaria osura an redue eelgrass shoot density. Aquat Bot 71, 149-154. http:// dx.doi.org/10.1016/s0304-3770(01)00183-8. Nelson TA, Nelson AV, Riarih H and Tjoelker M. 2003. Seasonal patterns in ulvoid algal iomass, produtivity, and key environmental fators in the Northeast Paifi. Bot Mar 46, 263-327. Nelson TA, Haerlin K, Nelson AV, Riarih H, Hothkiss R, Van Alstyne KL, Bukingham L, Simunds DJ and Fredrikson K. 2008. Eologial and physiologial ontrols of speies omposition in green maroalgal looms. Eology 89, 1287-1298. http://dx.doi.org/10.1890/07-0494.1. Niholls KH, Hopkins G, Standke SJ and Nakamoto L. 2001. Trends in total phosphorus in Canadian near-shore waters of the Laurentian Great Lakes: 1976-1999. J Gt Lakes Res 27, 402-422. http://dx.doi.org/10.1016/s0380-1330(01)70656-9. Nordy Ø and Hoxmark RC. 1972. Changes in ellular parameters during synhronous meiosis in Ulva mutailis Føyn. Exp Cell Res 75, 321-328. http://dx.doi.org/10.1016/0014-4827(72)90436-3. Nyerg CD and Wallentinus I. 2005. Can speies traits e used
664 나연주ㆍ전다빈ㆍ이정록ㆍ김영식ㆍ최한길ㆍ남기완 to predit marine maroalgal introdutions?. Biol Invasions 49, 688-694. http://dx.doi.org/10.1007/s10530-004-0738-z. Pekol P, Demeo-Anderson B, Rivers J, Valiela I, Maldonado M and Yates J. 1994. Growth, nutrient uptake apaities and tissue onstituents of the maroalgae Cladophora vagaunda and Grailaria tikvahiae related to site-speifi nitrogen loading rates. Mar Biol 121, 175-185. http://dx.doi. org/10.1007/bf00349487. Pitkänen H, Kondratyev S, Lääne A, Gran V, Kauppila P, Loigu E, Markovets I, Pahel K and Rumyantsev V. 1997. Pollution load on the Gulf of Finland from Estonia, Finland and Russia in 1985-1995. Summary report of the working group. In: Proeedings of the Final Seminar of the Gulf of Finland Year 1996, Environmental Institute, Helsinki, Finland, pp 9-18. Provasoli L. 1968. Media and prospets for the ultivation of marine algae. In: Cultures and Colletions of Algae. Watanae A. and Hattori A, eds. Proeeding of the US-Japan Conferene, Japanese Soiety for Plant Physiology, Tokyo, Japan, 63-75. Råerg S, Berger-Jönsson R, Björn A, Granéli E and Kautsky L. 2005. Effets of Pillayella littoralis on Fuus vesiulosus reruitment: impliations for ommunity omposition. Mar Eol Prog Ser 289, 131-139. http://dx.doi.org/10.3354/ meps289131. Serisawa Y, Yokohama Y, Aruga Y and Tanaka J. 2002. Growth of Eklonia ava (Laminariales, Phaeophyta) sporophytes transplanted to a loality with different temperature onditions. Phyol Res 50, 201 207. http://dx.doi.org/10.1046/ j.1440-1835.2002.00274.x. Shea R and Chopin T. 2007. Effets of germanium dioxide, an inhiitor of diatom growth, on the mirosopi laoratory ultivation stage of the kelp, Laminaria saharina. J Appl Phyol 19, 27-32. http://dx.doi.org/10.1007/s10811-006- 9107-x. Shimada S, Yokoyama N, Arai S and Hiraoka M. 2008. Phylogeography of the genus Ulva (Ulvophyeae, Chlorophyta), with speial referene to the Japanese freshwater and rakish taxa. J Appl Phyol 20, 979-989. http://dx.doi. org/10.1007/s10811-007-9296-y. Sokal RR and Rohlf FJ. 1995. Biometry, 3 rd Edition. Freeman, New York, U. S. A. Sousa AI, Martins I, Lilleø AI, Flindt MR and Pardal MA. 2007. Influene of salinity, nutrients and light on the germination and growth of Enteromorpha sp. spores. J Exp Mar Bio Eol 341: 142 150. http://dx.doi.org/10.1016/j. jeme.2006.09.020. Tan IH, Blomster J, Hansen G, Leskinen E, Maggs CA, Mann DG, Sluiman HJ and Stanhope MJ. 1999. Moleular phylogeneti evidene for a reversile morphogeneti swith ontrolling the gross morphology of two ommon genera of green seaweeds, Ulva and Enteromorpha. Mol Biol Eol 16, 1011-1018. Taylor R, Flether RL and Raven JA. 2001. Preliminary studies on the growth of seleted green tide algae in laoratory ulture: effets of irradiane, temperature, salinity and nutrients on growth rate. Bot Mar 44, 327-333. http://dx.doi. org/10.1515/bot.2001.042. Vahteri P, Mäkinen A, Salovius S and Vuorinen I. 2000. Are drifting algal mats onquering the ottom of the Arhipelago Sea, SW Finland?. Amio 29, 338-343. http://dx.doi. org/10.1579/0044-7447-29.6.338. Valiela I, MClelland J, Hauxwell J, Behr P, Hersh D and Foreman K. 1997. Maroalgal looms in shallow estuaries: ontrols and eophysiologial and eosystem onsequenes. Limnol Oeanogr 42, 1105-1118. Wang XH, Li L, Bao X and Zhao LD. 2009. Eonomi ost of an algae loom leanup in China s 2008 Olympi sailing venue. Eos Trans AGU 90, 238-239. http://dx.doi. org/10.1029/2009eo280002. Wang ZL, Xiao J, Fan SL, Li Y, Liu XQ and Liu DY. 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. Limnol Oeanogr 60, 1105-1117. http:// dx.doi.org/10.1002/lno.10083. Worm B, Heike K and Sommer U. 2001. Algal propagules anks modify ompetition, onsumer and resoure ontrol on Balti roky shores. Oeologia 128, 281-293. http:// dx.doi.org/10.1007/s004420100648. Ye NH, Zhang XW, Mao YZ, Liang CW, Xu D, Zhou J, Zhuang ZM and Wang QY. 2011. Green tides are overwhelming the oastline of our lue planet: taking the world s largest example. Eol Res 26, 477-485. http://dx.doi.org/10.1007/ s11284-011-0821-8. Zhang JH, Huo YZ, Yu KF, Chen QF, He Q, Han W, Chen LP, Cao JC, Shi DJ and He PM. 2013. Growth harateristis and reprodutive apaility of green tide algae in Rudong oast, China. J Appl Phyol 25, 795-803. http://dx.doi. org/10.1007/s10811-012-9972-4. Zhang JH, Huo YZ, Wu HL, Yu KF, Kim JK, Yarish C, Qin YT, Liu CC, Xu R and He PM. 2014. The origin of the Ulva maroalgal looms in the Yellow Sea in 2013. Mar Pollut Bull 89, 276-283. http://dx.doi.org/10.1016/j.marpolul.2014.09.049. Zhang JH, Kim JK, Yarish C and He P. 2016. The expansion of Ulva prolifera O.F. Müller maroalgal looms in the Yellow Sea, PR China, through asexual reprodution. Mar Pollut Bull 104, 101-106. http://dx.doi.org/10.1016/j.marpolul.2016.01.056. Zhao J, Jiang P, Liu Z, Wang J, Cui Y and Qin S. 2011. Geneti variation of Ulva (Enteromorpha) prolifera (Ulvales, Chlorophyta) the ausative speies of the green tides in the Yellow Sea, China. J Appl Phyol 23, 227-233. http://dx.doi. org/10.1007/s10811-010-9563-1.