한수지 49(5), , 2016 Original Article Korean J Fish Aquat Sci 49(5), ,2016 녹조대발생종금발대마디말 (Cladophora vadorum) 의절편생장에온도, 조도및영양염종류가미치는영향 나연주 전다

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1 한수지 49(5), , 2016 Original Artile Korean J Fish Aquat Si 49(5), ,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 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), Korean J Fish Aquat Si 49(5) , Otoer 2016 This is an Open Aess artile distriuted under the terms of the Creative Commons Attriution Non-Commerial Liens ( 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: Fax: address: hghoi@wku.a.kr Copyright 2016 The Korean Soiety of Fisheries and Aquati Siene 657 pissn: , eissn:

2 658 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)., (Cladophora vadorum)., Ha et al. (2016) (0.438 mg L -1 ) (0.046 mg L -1 ).,,. 재료및방법 (34 60' N, ' E) ,, (0.45 m)., (3 g) (100 ml) 8,000 rpm 10 s (Ø 0.5 mm) 3-4 (fragments). 성체의형태와발생, 6 ( 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,

3 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 ( 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, m (mean SE, n=50 ells), m (n=50) / m (Fig. 1A, 1B)., ( m, n=50) (Fig. 1C). 3,, 2 (Fig. 1D, 1E) mm 2 (Fig. 1F)., (Fig. 1C), (Fig. 1G). 8 (Fig. 1H, 1I). 온도와조도영향 m (mean SE, n=300 fragments), m. 1, m (50 mol photons m -2 s -1 ) 1, m (100 mol photons m -2 s -1 ) (F 1,16 =46.72, P<0.01). (15-30 ), m 25 2, m, 25 (P<0.01), (Tukey test, P 0.01, Tale 1) % day mol photons m -2 s -1 (Fig. 2). 영양염종류와생장, 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) < Irradiane (I) < Interation < 0.01 Error 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.

4 660 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 ) μmol photons m -2 s 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) m % day M (100 M) (Fig. 3A)., 3 10, 20, 100 M (Tukey HSD test, P 0.01) M, , m % day -1 (Fig. 3B). 80 M 100 M (F 5,12 =104.49, P<0.01) m % day -1 (Fig. 3C). (0, 5, 10, 40 M) (80, 100 M) (F 6,14 =99.13, P<0.01), 200 M M,, (0, 5 M) 10, M (Tukey test, P 0.01).

5 녹조대발생종금발대마디말성체의절편생장 661 (A) (B) (C) RGR (% day -1 ) RGR (% day -1 ) RGR (% day -1 ) a a a PO 4 (μmol/l) a NH 4 (μmol/l) 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) , , (Cladophora alida) (C. vagaunda) 25 Breeman et al. (2002). (50 mol photons m -2 s -1 ) 100 mol photons m -2 s -1, ( mol photons m -2 s -1 )., % day % day -1 (16-26, 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) ( ) 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) ( 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 M 10-15% day -1 (de Paula Silva et al., 2008).

6 662 나연주ㆍ전다빈ㆍ이정록ㆍ김영식ㆍ최한길ㆍ남기완 15-17% day -1 (4-8% day -1 ) Cladophora oelothrix (10-15% day -1 ) ( 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) , 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 Exessive growth of Cladophora (Dillwyn) kützing and enteri ateria in mats in the Southwestern Istanul oast, Sea of Marmama. IUFS J Biol 72, Blomster J, Bäk S, Fewer DP, Kiirikki M, Lehvo A, Maggs CA and Stanhope MJ Novel morphology in Enteromorpha (Ulvophyeae) forming green tides. Am J Bot 89, Boudouresque CF and Verlaque M Biologial pollution in the Mediterranean Sea: invasive versus introdued marophytes. Mar Pollut Bull 44, S X(01) Breeman A, Oh YS, Hwang MS and Van Den Hoek C Evolution of temperature responses in the Cladophora vagaunda omplex and the C. alida/seriea omplex (Chlorophyta). Eur J Phyol 37, /S Charlier RH, Morand P, Finkl CW and Thys A Green tides on the Brittany oasts. Environ Res Eng Manage 3, Choi TS, Kang EJ, Kim JH and Kim KY Effets of salinity on growth and nutrient uptake of Ulva pertusa (Chlorophyta) from an eelgrass ed. Algae 25, dx.doi.org/ /algae Dan A, Ohno M and Matuoka M 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 Integrating filamentous green tide algae into tropial pondased aquaulture. Aquaulture 284, org/ /j.aquaulture Deng YY, Tang XR, Huang BX and Ding LP Life history of Chaetomorpha valida (Cladophoraeae, Chlorophyta) in ulture. Bot Mar 54, BOT Dodds WK and Gudder DA The eology of Cladophora. J Phyol 28, Duarte C Sumerged aquati vegetation in relation to different nutrient regimes. Ophelia 41, g/ / Eaton JW, Brown JG and Rouno FE Some oservations on polarity and regeneration in Enteromorpha. Br Phyol Bull 3, Entwisle TJ Phenology of the Cladophora-Stigeolonium ommunity in two uran reeks of Melourne. Aust J Mar Freshwater Res 40, MF Flether RL The British Isles. In: Sharmm W, Nienhuis PH(eds) Marine enthi vegetion: reent hanges, the ef-

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한수지 49(2), , 2016 Note Korean J Fish Aquat Sci 49(2), ,2016 부안상록해수욕장의사상녹조류금발대마디말 (Cladophora vadorum) 대량발생과영양염농도 하동수 유현일 장수정 1 황은경 * 국립수산

한수지 49(2), , 2016 Note Korean J Fish Aquat Sci 49(2), ,2016 부안상록해수욕장의사상녹조류금발대마디말 (Cladophora vadorum) 대량발생과영양염농도 하동수 유현일 장수정 1 황은경 * 국립수산 한수지 49(2), 241-246, 2016 Note Korean J Fish Aquat Sci 49(2),241-246,2016 부안상록해수욕장의사상녹조류금발대마디말 (Cladophora vadorum) 대량발생과영양염농도 하동수 유현일 장수정 1 황은경 * 국립수산과학원해조류연구센터, 1 국립수산과학원서해수산연구소자원환경과 Bloom of a Filamentous