쇠미역과미역배우체의생장과성숙 191 Park et al. (11) 2 (, mm;, mm), (12, 17, 22 ) (2, 1,, 6, 1, 18 µmol photons m -2 s -1 ) 22., (17, 6 µmol photons m
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1 한수지 49(2), , 16 Original Article Korean J Fish Aquat Sci 49(2),19-197,16 쇠미역 (Costaria costata) 과미역 (Undaria pinnatifida) 배우체의생장과성숙에광과온도가미치는영향 나연주 전다빈 한수진 Cyr Abel Ogandaga Maranguy 안대성 1 차형기 2 이재봉 3 양재형 4 이해원 5 최한길 * 원광대학교생명과학부 / 기초자연과학연구소, 1 생태와환경, 2 국립수산과학원연근해자원과, 3 국립수산과학원원양자원과, 4 동해수산연구소자원환경과, 5 국립수산과학원연구협력과 Crossed Effects of Light and Temperature on the Growth and Maturation of Gametophytes in Costaria costata and Undaria pinnatifida Yeon Ju Na, Da Vine Jeon, Su Jin Han, Cyr Abel Ogandaga Maranguy, Dae Sung An 1, Hyung Kee Cha 2, Jae Bong Lee 3, Jae Hyeong Yang 4, Hae Won Lee 5 and Han Gil Choi* Faculty of Biological Science and Institute for Basic Science,Wonkwang University, Iksan 54538, Korea 1 Institute of Eco and Environmental Incorporation, Bucheon 1451, Korea 2 Coastal Water Fisheries Resources Research Division, National Institute of Fisheries Science, Busan 4683, Korea 3 Distant Water Fisheries Resources Research Division, National Institute of Fisheries Science, Busan 4683, Korea 4 Fisheries Resources and Environment Research Division, National Institute of Fisheries Science, Gangneung 25435, Korea 5 Research Cooperation Division, National Institute of Fisheries Science, Busan 4683, Korea We examined for differences in the growth and fertility of microscopic stages between Costaria costata and Undaria pinnatifida in crossed temperature irradiance (12, 17, 22, 6 μmol photons m 2 s 1 ) and daylength irradiance (8, 12, 16, 24 h, 6 μmol photons m 2 s 1 ) experimental designs. After 13 days in culture, maximum gametophyte growths of the two species were obtained under the following combinations of factors: 17 6 μmol photons m 2 s 1 and 24 h daylength 6 μmol photons m 2 s 1. C. costata gametophytes produced sporophytes about 6 days earlier than those of U. pinnatifida. Sporophyte densities were highest in the 12 h daylength 6 μmol photons m 2 s 1 combination after days in C. costata (9.7 plants mm 2 ) and 26 days in U. pinnatifida (9.1 plants mm 2 ). However, optimal growth conditions for the microscopic sporophytes differed between species: 17 6 μmol photons m 2 s 1 for C. costata, 12 6 μmol photons m 2 s 1 for U. pinnatifida. Thus, C. costata gametophytes grew faster and produced sporophytes earlier than those of U. pinnatifida, even though the haploid gametophytes of the two species responded similarly to environmental conditions. Optimal growth temperatures for the microscopic sporophytes also differed between species. Key words: Costaria costata, Fertility, Growth, Gametophyte, Undaria pinnatifida 서론 (Costaria costata) (Undaria pinnatifida) (Laminariales) (Choi et al., 7)., (blade) (Cho, 1), (midrib), (stipe) (sporophyll) (Alariaceae). (Sohn, 4). Korean J Fish Aquat Sci 49(2) , April 16 This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial Licens ( which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. Received 26 February 16; Revised 21 April 16; Accepted 25 April 16 *Corresponding author: Tel: Fax: address: hgchoi@wku.ac.kr Copyright 16 The Korean Society of Fisheries and Aquatic Science 19 pissn: , eissn:
2 쇠미역과미역배우체의생장과성숙 191 Park et al. (11) 2 (, mm;, mm), (12, 17, 22 ) (2, 1,, 6, 1, 18 µmol photons m -2 s -1 ) 22., (17, 6 µmol photons m -2 s -1 ) (12 h, 35 psu) (15 psu). (,,,,, ) (Kim and Nam, 1997; Morita et al., 3; Wu et al., 4; Choi et al., 5; Morelissen et al., 13),., 1962, ( ) ( ) (78 ) 25.66% ( 13.93%) 3 (, go.kr)., 5-1 m, (Kim et al., 6)., (strain),, (Wu et al., 4; Wi et al., 8; Hwang et al., 1). (hybrid) (Pang et al., 1997; Morita et al., 3; Wu et al., 4; Choi et al., 5; Druehl et al., 5),,,, (Wu et al., 4; Westermeier et al., 6; Hwang et al., 9; Xu et al., 9; Hwang et al., 1). (Wu et al., 1994; Kaas, 1998; Li et al., 1999; Wu et al., 4; Xu et al., 9), (Xu et al., 9).,,,. 재료및방법 (34 28' N, ' E) 12 5 (5-1 )... ( cm) Petri dish (Ø 9 cm).1 ml 17 6 µmol photons m -2 s h. PESI (Tatewaki, 1966) 3 ml Petri dish (Ø 1 cm) 3 12 h (12, 17, 22, 27 ) (, 6 µmol photons m -2 s -1 ), 17 4 (8, 12, 16, 24 h) 2 (, 6 µmol photons m -2 s -1 ) 3 13., (DX-). 3 GeO 2 (5 mgl -1 ) (Shea and Chopin, 7). Image J. (RGR, relative growth rate), 13 (Serisawa et al., 2). RGR (% day -1 ) = 1 ln (L 2 - L 1 ) / T 2 - T 1 L 2 13, L 1 (embryospore), T 2 -T (12, 17, 22 ) (, 6 µmol photons m -2 s -1 ) (8, 12 h) (, 6 µmol photons m -2 s -1 ). (egg), 26. Petri dish (Ø 9 cm) 4 mm 2 cover glass
3 192 나연주ㆍ전다빈ㆍ한수진ㆍ Cyr Abel Ogandaga Maranguy ㆍ안대성ㆍ차형기ㆍ이재봉ㆍ양재형ㆍ이해원ㆍ최한길., 26, 12 h, 17 6 µmol photons m -2 s , Cochran s test (homogeneity of variances), two-way ANOVA test., Tukey s HSD test (Sokal and Rohlf, 1995). STATIS- TICA version 5.. 배우체생장 결과 3 (12, 17, 22 ) 2 (, 6 µmol photons m -2 s -1 ), 27, 2 (Fig. 1). 13, m (RGR, % day -1 ) 17 (F 3,16 =972.1, P<.1), µmol photons m -2 s -1 6 µmol photons m -2 s -1 (F 1,16 =46.68, P<.1)., (F 3,16 =6., P<.1), (Tukey HSD test). 13, m (RGR, %day -1 ) 17 (F 3,16 = , P<.1), µmol photons m -2 s -1 6 µmol photons m -2 s -1 (F 1,16 =15.78, P<.1)., (F 3,16 =.26, P<.1), (Tukey HSD test). (8, 12, 16, 24 h) (, 6 µmol photons m -2 s -1 ) µm ( %day -1 ) (24 h) (F 3,16 =265.1, P<.1), µmol photons m -2 s -1 6 µmol photons m -2 s -1 (F 1,16 =52.15, P<.1, Fig. 2)., (F 3,16 =1.24, 5 Female length (µm) µmol photons m -2 s * (A) Female length (µm) µmol photons m -2 s (A) Female length (µm) µmol photons m -2 s -1 (B) Female length (µm) µmol photons m -2 s -1 (B) Temperature( ) * Daylength (h) Fig. 1. Average female lengths of (A) Costaria costata and (B) Undaria pinnatifida gametophytes cultured for 13 days under various temperature and irradiance levels. Daylength was constant as 12h. Values represent mean ±SE (n=3 replicates). *, no data Fig. 2. Average female lengths of (A) Costaria costata and (B) Undaria pinnatifida gametophytes cultured for 13 days under four daylength and two irradiance levels. Temperature was constant as 17. Values represent mean ±SE (n=3 replicates).
4 쇠미역과미역배우체의생장과성숙 193 Length / width (µm) Costaria costata Length Species Width Undaria pinatifida Fig. 3. Average cell lengths and widths of Costaria costata and Undaria pinnatifida female gametophytes having young sporophytes after 26 days in the same culture conditions of 12 h daylength, 17, and 6 µmol photons m -2 s -1. P =.33), (Tukey HSD test) µm ( %day -1 ) (24 h) (F 3,16 =197.86, P<.1), ( µmol photons m -2 s -1 ) (6 µmol photons m -2 s -1 ) (F 1,16 =28.57, P<.1)., (F 3,16 =2.36, P =.11), (Tukey HSD test) µmol photons m -2 s -1, 22. 포자체형성 2 ( ) 26 ( ). (12, 17, 22 ) (, 6 µmol photons m -2 s -1 ) /mm µmol photons m -2 s -1 (Table 1) / mm µmol photons m -2 s -1. (8, 12, 16, 24 h) (, 6 µmol photons m -2 s -1 ) 12 h 6 µmol photons m -2 s /mm 2, 12 h (16, 24 h) (Table 1)., 12 h 6 µmol photons m -2 s / mm 2. 포자체생장 µm 17 6 µmol photons m -2 s -1 Table 1. Average density (plants/mm 2 ) of young sporophytes that produced from Costaria costata and Undaria pinnatifida gametophytes grown under various temperature, daylength, and irradiance levels. Density was estimated after and 26 days for Costaria costata and Undaria pinnatifida, respectively. Values represent mean ±SE (n=3 replicates) Species C. costata U. pinnatifida Irradiance Temperature ( ) Daylength (h) (µmol photons m -2 s -1 ) ± ± ± ±.5 8.5± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ±.55 Table 2. Average lengths(µm) of young sporophytes produced from Costaria costata and Undaria pinnatifida gametophytes grown under various temperature, daylength, and irradiance levels. Growth was measured after and 26 days for Costaria costata and Undaria pinnatifida, respectively. Values represent mean ±SE (n=3 replicates) Species C. costata U. pinnatifida Irradiance Temperature ( ) Daylength (h) (µmol photons m -2 s -1 ) ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ±3.84
5 194 나연주ㆍ전다빈ㆍ한수진ㆍ Cyr Abel Ogandaga Maranguy ㆍ안대성ㆍ차형기ㆍ이재봉ㆍ양재형ㆍ이해원ㆍ최한길 µm 12 6 µmol photons m -2 s -1 (Table 2). (F 3,16 = , P<.1, F 3,16 =485.75, P<.1) (F 1,16 =82.51, P<.1, F 1,16 = 98.23, P<.1) µm 6 µmol photons m -2 s h µm (6 µmol photons m -2 s -1, 12 h) (Table 2). (F 1,16 =113.49, P<.1, F 1,16 =6.57, P<.1) (F 3,16 = , P<.1, F 3,16 = , P<.1). 26, 2 12 h, 17 6 µmol photons m -2 s µm (n=4 cells) µm (n=4), µm (n=4 cells) µm(n=4) 고찰,, (Choi et al., 8; Oh et al., 15)., (, ) (Kim et al., 5; Kim et al., 7a; Choi et al., 8), ( ) (Saito, 1975).,, (Akiyama, 1965; Morita et al., 3; Wu et al., 4; Choi et al., 5), (16 h) (24 h) (Tseng et al., 1959; Akiyama, 1965; Lüning, 1981; Choi et al., 5)., 17-, (Saito, 1956; Kang and Koh, 1977), Kim and Nam (1997) 12 h (5, 17,, 25, 3 ) (, 4, 6, 1 µmol photons m -2 s -1 ) 17 6 µmol photons m -2 s µmol photons m -2 s -1 (Wi et al., 8; Hwang et al., 1), 15 (Hwang et al., 1) 25 (Kim et al., 7a), ( ), (5-6 ) (Table 3)., Wu et al. (4) (, ) 5-22, 25 (12:12h LD), 37.1 %day -1 (8 µmol photons m -2 s -1 ) 38.6 %day -1 (1 µmol photons m -2 s -1 ). 2 ( ) 17 6 µmol photons m -2 s -1, 13 (36.6% day -1 ) (33.91 %day -1 )., (12 h) (8 h). (16 h), 12 h (Choi et al., 5). Liu et al. (4) (egg-discharge) 11h (11:13h LD) 5-15, (24 h). (1, 12, 14 h) (Wi et al., 8), (Hwang et al., 1). Tseng et al. (1959), Lüning (1981).,,. 17, 6 µmol photons m -2 s -1, 12 h, (17 ) (12 ). Saito (1975) 15-17, (5-1 mm) 17 (Park et al., 11),,., 1
6 쇠미역과미역배우체의생장과성숙 195 Table 3. Optimal growth conditions in the microscopic gametophytes and sporophytes of various kelp species growing around the coastal areas of Korea. The tested environmental conditions represents in the parenthesis Species Gametophyte Costaria costata Undaria pinnatifida Ecklonia cava Ecklonia stolonifera Ecklonia stolonifera Saccharina sculpera Saccharina sculpera Temperature ( ) 17 (12, 17, 22, 27) 17 (12, 17, 22, 27) 15 (5, 1, 15, ) 15 (5,1,15, ) 25 (5,1,15,,25) (5,1,15,,25) (5,1,15,,25) Saccharina japonica (5,1,15, ) Sporophyte Costaria costata Undaria pinnatifida Ecklonia stolonifera Saccharina sculpera Saccharina japonica Saccharina japonica ND; no data. 17 (12, 17, 22) 12 (12, 17, 22) (1,15,,25) 1 (5,1,15) 1 (5,1,15) 1 (5,1,15, ) Irradiance (µmol photons m -2 s -1 ) 6 (, 6) 6 (, 6) (5, 1,, 4) (5, 1,, 4) (, 4, 6, 8) 4 (, 4, 6, 8) 4 (4, 8, 1) 1 (1, 3, 7, 15) 6 (, 6) 6 (, 6) (, 4, 6, 8) 4 (4, 8, 1) 8 (4, 8, 1) 7 (1, 3, 7) Daylength (h) 24 (8, 12, 16, 24) 24 (8, 12, 16, 24) 14 (1, 12, 14) 12 (1, 12, 14) ND ND References This study This study Wi et al., 8 Hwang et al., 1 Kim et al., 7a Kim et al., 7b ND Kim et al., 5 ND Kang and Koh, (8, 12) 12 (8, 12) ND This study This study Kim et al., 7a ND Kim et al., 5 ND Kim et al., 5 ND Kang and Koh, 1999, ( 25, ) (Table 3).,.,, 2,. 2 (16 h) (24 h) (8 h) (12 h).,,. 사사 15 (R163)., 15.
7 196 나연주ㆍ전다빈ㆍ한수진ㆍ Cyr Abel Ogandaga Maranguy ㆍ안대성ㆍ차형기ㆍ이재봉ㆍ양재형ㆍ이해원ㆍ최한길 References Akiyama K Studies of ecology and culture of Undaria pinnatifida (Harv.) Sur. II. Environmental factors affecting the growing and maturation of gametophyte. Bull Tohoku Reg Fish Res Lab 25, Cho GY 1. Laminariales. In: Algal Floral of Korea. Volume 2, Number 2. Heterokontophyta: Phaeophyceae; Ishigeales, Dictyotales, Desmarestiales, Sphacelariales, Cutleriales, Ralfisales, Laminariales. Anon., eds. National Institute of Biological Resources, Incheon, Korea, Choi HG, Kim YS, Lee SJ, Park EJ and Nam KW. 5. Effects of daylength, irradiance and settlement density on the growth and reproduction of Undaria pinnatifida gametophytes. J Appl Phycol 17, Choi HG, Kim YS, Lee SJ and Nam KW. 7. Growth and reproductive patterns of Undaria pinnatifida sporophytes in a cultivation farm in Busan, Korea. J Appl Phycol 19, Choi HG, Lee KH, Yoo HI, Kang PJ, Kim YS and Nam KW. 8. Physiological differences in the growth of Sargassum horneri between the germling and adult stages. J Appl Phycol, Druehl LD, Collins JD, Lane CE and Saunders GW. 5. An evaluation of methods used to assess intergeneric hybridization in kelp using pacific Laminariales (Phaeophyceae). J Phycol 41, Hwang EK, Baek JM and Park CS. 9. The mass cultivation of Ecklonia stolonifera Okamura as a summer feed for the abalone industry in Korea. J Appl Phycol 21, Hwang EK, Gong YG, Ha DS and Park CS. 1. Inducing the regeneration and maturation of free-living gametophytes of Ecklonia stolonifera Okamura (Laminariales, Phaeophyta). Kor J Fish Aquat Sci 43, kfas Kaas R The seaweed resources of France. In: Seaweed Resources of the World. Critchley AT and Ohno M, eds. Japan International Cooperation Agency, Yokosuka, Japan, Kang JW and Koh NP Seaweed Culture. Taewha press, Pusan, Korea, Kang RS and Koh CH Germination and growth of Laminaria japonica (Phaeophyta) microscopic stages under different temperature and photon irradiances. J Kor Fish Soc 32, Kim YD, Song HI, Hong JP, Jeon CY, Kim SK, Han HK, Kim DS and Bang JK. 6. Growth and maturation of the brown seaweed Costaria costata transplanted for the wildstock enhancement. J Life Sci 16, org/1.5352/jls Kim DS, Hong JP, Kim YD, Song HI and Kim HG. 7a. Seed production and cultivation of Ecklonia stolonifera Okamura, Phaeophyta. J Aquacult, Kim JH, Kim YD, Song HI, Gong YG, Lee SB and Jin HJ. 7b. Separation and preservation of the male and female gametophytes of Kjellmaniella crassifolia Miyabe (Phaeophyta). J Kor Fish Soc 4, org/1.5657/kfas Kim HG, Park JG and Kim DS. 5. Comparative laboratory culture studies of the native kelp Kjellmaniella crassifolia and the introduced kelp Laminaria japonica in east coast of Korea. J Aquacult 18, Kim YS and Nam KW Temperature and light responses on the growth and maturation of gametophytes of Undaria pinnatifida (Harvey) Suringar in Korea. J Kor Fish Soc 3, Li DP, Zhou ZG, Liu HH and Wu CY A new method of Laminaria japonica strain selection and seedling culture by the use of gametophyte clones. Hydrobiologia 398/399, Liu J, Dong L, Shen Y and Wu C. 4. Effect of light period on egg-discharge of gametophyte clones of Undaria pinnatifida (Phaeophyta). J Appl Phycol 16, Lüning K Egg release in gametophytes of Laminaria saccharina: Induction by darkness and inhibition by blue light and UV. Br Phycol J 16, org/1.18/ Morita T. Kurashima A and Maegawa M. 3. Temperature requirements for the growth and maturation of the gametophyte of Undaria pinnatifida and U. undarioides (Laminariales, Phaeophyceae). Phycol Res 51, org/1.146/j t x Morelissen B, Dudley BD, Geange SW and Phillips NE. 13. Gametophyte reproduction and development of Undaria pinnatifida under varied nutrient and irradiance conditions. J Exp Mar Biol Ecol 448, Oh JC, Yoo OH and Choi HG. 15. Interactive effects of increased temperature and pco 2 concentration on the growth of a brown algae Ecklonia cava in the sporophyte and gametophyte stages. Ocean Polar Res 37, 1-9. Pang SJ, Hu XY, Wu CY, Hirosawa A and Ohno M Intraspecific crossings of Undaria pinnatifida (Harv.) Sur. A possible time-saving way of strain selection. Chin J Oceanol Limnol 15, Park SK, Heo JS, Kim BY, Song JN, Lim GY, Kim HN and Choi HG. 11. Comparison on the growth of Costaria costata and Undaria pinnatifida sporophytes in culture and their field populations. Kor J Fish Aquat Sci 44, Saito Y An ecological study of Undaria pinnatifida Sur.
8 II. On the influence of environmental factors upon the maturity of gametophytes and early development of sporophytes. Bull Jpn Soc Sci Fish 22, org/1.2331/suisan Saito Y Practical significance of algae in Japan. In: Advance phycology in Japan. Tokida J and Hirose H. eds. VEB Gustav Fischer Verlag, Jena, Netherlands, Serisawa Y, Yokohama Y, Aruga Y and Tanaka J. 2. Growth of Ecklonia cava (Laminariales, Phaeophyta) sporophytes transplanted to a locality with different temperature conditions. Phycol Res 5, j x Shea R and Chopin T. 7. Effects of germanium dioxide, and inhibitor of diatom growth, on the microscopic laboratory cultivation stage of kelp, Laminaria saccharina. J Appl Phycol 19, Sohn CH. 4. Developments on two newly cultivated species Capsosiphon fulvescens and Costaria costata in Korea. J Phycol Suppl 39, Sokal RR and Rohlf FJ Biometry, 3rd edn. WH Freeman. New York. USA. Tseng CK, Ren KZ and Wu CY On the discharge of eggs and spermatozoids of Laminaria japonica and the morphology of the spermatozoids. Kexue Tongbao 4, Tatewaki M Formation a crustaceous sporophyte with unilocular sporangia in Scytosiphon lomentaria. Phycologia 6, Westermeier R, Patiño D, Piel MI, Maier I and Mueller DG. 6. A new approach to kelp mariculture in Chile: production of free-floating sporophyte seedlings from gametophyte cultures of Lessonia trabeculata and Macrocystis pyrifera. Aqua Res 37, Wi MY, Hwang EK, Kim SC, Hwang MS, Baek JM and Park CS. 8. Regeneration and maturation induction for the free- living gametophytes of Ecklonia cava Kjellman (Laminariales, Phaeophyta). J Kor Fish Soc 41, dx.doi.org/1.5657/kfas Wu CY, Pang SJ and Jiang BY Establishment and application of vegetative gametophytes of Laminaria japonica. In: Seaweed Biotechnology. Wang SJ, ed. Shanghai Scientific and Technical Publishers, Shanghai, China, Wu CY, Li DP, Liu HH, Peng GA and Liu JX. 4. Mass culture of Undaria gametophyte clones and their use in sporeling culture. Hydrobiologia 512, Xu B, Zhang QS, Qu SC, Cong YZ and Tang XX. 9. Introduction of a seedling production method using vegetative gametophytes to the commercial farming of Laminaria in China. J Appl Phycol 21, s z 쇠미역과미역배우체의생장과성숙 197
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