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1 한국어병학회지제26권제3호 (2013) J. Fish Pathol., 26(3) : 265~274 한국산진두발, Chondrus ocellatus 의생장과질병에대한연구 이순정 박명애 아벨 * 박서경 * 김회경 ** 김영식 *** 최한길 * 13) 국립수산과학원수산생물방역과, * 원광대학교생명과학부, ** 원광대학교식품영양학과, *** 군산대학교해양생물공학과 A Study on the Growth and Disease of Chondrus ocellatus in Korea Soon Jeong Lee, Myoung-Ae Park, Cyr Abel Ogandaga-Maranguy *, Seo Kyoung Park *, Hoikyung Kim **, Young Sik Kim *** and Han Gil Choi * Aquatic Life Disease Control Division, NFRDI, Busan , Korea * Faculty of Biological Science, Wonkwang University, Iksan, Jeonbuk , Korea, ** Department of Food and Nutrition, Wonkwang University, Iksan, Jeonbuk , Korea, *** Department of Marine Biotechnology, Kunsan National University, Kunsan , Korea The growth, reproduction, gametophyte(g)/tetrasporophyte(t) ratio, and diseases of Chondrus ocellatus populations were examined at 3 sites (Samcheok, Youngduk, Pohang) of East coast and at 2 sites (Hakampo, Manripo) of West coast in between July and August, Average plant lengths were cm and it was minimum at Manripo and maximum at Pohang population. In general, average plant length and weight of C. ocellatus were greater on East coast than West coast populations. The proportion of vegetative plant was between %, and G/T ratio of total plants including vegetative plants after testing resorcinol method was 3:2 on the East coast where is gametophyte dominant area. However, G/T ratio was 1:1 at Hakampo and 1:2.3 at Manripo representing tetrasporophyte dominance. In the present study, Korean C. ocellatus have various diseases (white colour and green colour), an endophytic alga(ulvella sp.), and many epiphytic macroalgae and diatoms. Healthy C. ocellatus plants were about % in summer population and most of plants had disease. Chondrus ocellatus had a filamentous green alga, endophytic Ulvella sp. which was not identified. The endophyte is easily observed in C. crispus growing in Europe and Canada and it is recognised as a pathogen destroying population and reducing yield of C. crispus. Thus more interest and research on the endophytic algae and disease of C. ocellatus are required. Key words : Chondrus ocellatus, Disease, Endophyte, Growth 해조류는일차생산자, 생물지표자, 생물여과자의생태학적기능을가지며인간에게는식용, 의약품, 화장품, 해조산업과바이오에탄올의원료로서관심이증대하고있다 (Vasquez and Guerra, 1996; Orfanidis Corresponding author: Han Gil Choi Tel: , Fax: hgchoi@wku.ac.kr et al., 2001; Arévalo et al., 2007; Tribollet and Vroom, 2007). 하지만, 인간활동으로인한환경오염과지구온난화에의한수온상승, 해양산성화및해조류질병증가는연안역에서해조류종다양성과생물량감소를초래하고있다 (Orfanidis et al., 2001; Arévalo et al., 2007).

2 266 이순정 박명애 아벨 박서경 김회경 김영식 최한길 해조류질병연구는 Andrews(1976) 에의해최초로시작되었으며, 이후다양한분류군에서병리연구 (Correa and Flores, 1995; Largo et al., 1995; Craigie and Correa, 1996; Largo et al., 1998) 가수행되었다. 질병원인균에대한분자생물학적동정과정량적연구등으로전문화되고있으나다른수산생물에비해초보적수준이다 (Narita et al., 2001; Park et al., 2001a; 2001b; 2006; Hattori et al., 2007). 또한, 해조류질병연구는한국, 중국과일본의연구자에의해양식대상종인김 (Porphyra spp.), 미역 (Undaria pinnatifida), 꼬시래기류 (Gracilaria spp.) 와다시마류 (Laminaria spp.) 에서주로이루어졌으나, 최근에는동남아시아에서상업종인 Kappaphycus spp. 와 Eucheuma의 ice-ice질병에대한연구가증가하는추세이다 (Amano et al., 1995; Largo, 2002; Butardo et al., 2003). 우리나라에서해조류질병에관한연구는김, 미역, 다시마와같은양식종에한정되어수행되었으나정보가부족한실정이다 (Kang and Koh, 1977; Park et al., 2009). 해조류의질병은양식대상종의수확량감소와상품성하락등어업인의소득감소와직결되기때문에많은정보와연구자가시급하게요구되는상황이다. 진두발 (Chondrus) 속 (genus) 은동형삼상세대교번을하는다년생홍조류로서온대와냉온대해역에넓게분포하고있으며 (Lüning, 1985), 젤리와음식물의안정제인카라기난 (carrageenans) 의원료를생산하는해조류로서, 상업적인중요성때문에생리 생태학적연구가많이수행되었다 (Chopin and Floch, 1987; Bates and Craigie, 1988; Taylor and Chen, 1994; Garbary et al., 2011). 유럽에서 Irish moss 라고불리는주름진두발 (Chondrus crispus) 은북대서양에널리분포하며, 반수체 (haploid) 인암 수배우체는 kappa (κ)-와 iota-카라기난을합성하고사분포자체 ( 배수체, diploid) 는 lambda(λ)-카라기난을생산하여생활사에따라생화학적물질이다르게나타났다 (McCandless et al., 1973; Chopin et al., 1987; Zablackis et al., 1991; Garbary et al., 2011; McLachlan et al., 2011). 배우체와포자체의카라기난성분이다른점을이용한 acetal-resorcinol 실험으로진두발의영양체 (vegetative plant) 를사분포자체와배우체로구분하여개체군연구에이용되고있다 (Craigie and Leigh, 1978). Bouarab et al. (2001) 은 C. crispus를숙주로서식하는내생조류인녹조류 Acrochaete operculata 는 λ-카라기난을함유한사분포자체에쉽게감염되며, κ-카라기난을가지는배우체는저항성이있다고하였다. 프랑스의해조산업에서카라기난원료로서중요한 C. crispus 배우체의야외개체군은과도한수확으로지속적으로감소하여 (Braud and Delepine, 1981), 빠른생장을보이는배우체선택연구 (Guiry, 1981) 와원형질체생산을위한연구가수행되었다 (Gall et al., 1990). 진두발 (Chondrus ocellatus) 은한국, 일본, 중국과대만해안에주로분포하며 (Tayler and Chen, 1994), 생활사, 형태와생장에대한연구가일부수행되었다 (Ji and Guo, 1992; Brodie et al., 1993; Li et al., 1994). 한국에서진두발은파도노출이중간정도인해안의조간대하부에번무하며 (Kang, 1968), 진두발의야외개체군의생장과성숙에관한생물계절학적연구 (Kim et al., 2006) 와진두발추출물은항혈액응고성분을가진다는연구가있다 (Choi and Kim, 1999). 따라서, 본연구는해조콜로이드산업의중요한부분을차지하는카라기난을생산하는원료해조류인진두발의미래양식을위한정보를제공하기위하여, 1) 우리나라동해안과서해안에서식하는진두발야외개체군의여름철생장비교, 2) 종묘생산을위한진두발의성숙비율, 3) 서로다른카라기난을가지는사분포자체와배우체비율비교, 4) 진두발의질병현황과빈도에대한자료를확보할목적으로수행되었다.

한국산진두발, Chondrus ocellatus 의생장과질병에대한연구 267 재료및방법본연구를위해진두발은동해안 3개정점 ( 삼척, 포항, 영덕 ) 과서해안의 2개정점 ( 학암포와만리포 ) 의조간대하부에서 2013년 7-8월에간조시에조체별로 50 cm 이상의간격을두고채집하였다 (Fig. 1).

3 한국산진두발, Chondrus ocellatus 의생장과질병에대한연구 267 재료및방법본연구를위해진두발은동해안 3개정점 ( 삼척, 포항, 영덕 ) 과서해안의 2개정점 ( 학암포와만리포 ) 의조간대하부에서 2013년 7-8월에간조시에조체별로 50 cm 이상의간격을두고채집하였다 (Fig. 1). 질병현황진두발개체군별로 50개체를무작위로선택하여육안으로외부형태를관찰하여질병유무를확인하였고해부현미경하에서질병의심개체를관찰하고질병별개체수를계수하였다. 내생조류 (endophyte) 는면도날로진두발조직을얇게절편한후광학현미경으로관찰하였고사진촬영을실시하였다. 또한, 진두발의착생해조류 (epiphyte) 는현미경하에서동정하였으며, 출현종의학명및국명은 Lee and Kang (2002) 에따랐다. 결과 Fig. 1. Collection sites of Chondrus ocellatus for this study. 생장과생식상태정점별로진두발 30개체를무작위로선택하여조체별길이와무게 ( 습중량 ) 를측정한후, 생식상태를낭과를가진과포자체, 사분포자낭을사분포자체와생식기관이발달하지않은영양체로구분하였다. 야외개체군에서다른형태의카라기난을함유한배우체 ( 암 수 ) 와포자체의비율이정점에따라다르게나타나는지를확인하기위하여영양체로부터확보한원형디스크 ( 지름 2 cm) 의수분을제거하고 Lazo et al. (1989) 의방법에따라 acetal-resorcinol test를실시하여 κ-카라기난을포함하고있어서붉은색 (red) 과 λ-카라기난을세포벽에가짐으로써분홍색 (pink) 을보이는사분포자체로분류하였다. 생장여름철진두발의정점별평균체장은 cm (n=30 plants) 로서만리포에서최소였고포항에서최대길이생장을보였다 (Fig. 2A). 진두발의최대체장은동해안의삼척에서 cm, 영덕에서 cm, 포항에서 cm였으며, 서해안의학암포에서 cm, 만리포에서 cm로측정되어동해안개체가서해안진두발에비해큰것을확인하였다. 진두발의평균무게 ( 습중량 ) 는정점별로 g (n=30 plants) 이었으며 (Fig. 2B) 만리포에서최소였고영덕에서최대였다. 금번에조사한 150개개체에서가장작은개체는 0.03 g이었고최대습중량은영덕에서채집된개체로 g을나타냈다. 동해안진두발의평균습중량은 g (n=30) 로서서해안학암포의 2.45 g과만리포의 1.48에비해최대 4배이상높은것으로확인되었다 (Fig. 2). 생식상태동해안진두발개체군의영양체의비율은 % 였으며 (Table 1), 서해안의만리포는 26.7 %, 그리고학암포는 40.0 % 로서정점별차이를보였으며, 사분포자체는동해한포항에서최대 33.3 % 였으

4 268 이순정 박명애 아벨 박서경 김회경 김영식 최한길 Table 1. The number of plant and percentage(%) of reproductive plants in Chondrus ocellatus populations collected on the rocky shores of East and West coasts, Korea Reproduction East Sea West Sea Samcheok Youngduk Pohang Hakampo Manripo Tetrasporic 7(23.3%) 7(23.3%) 10(33.3%) 11(36.7%) 14(46.7%) Caposporic 12(40.0%) 3(10.0%) 10(33.3%) 7(23.3%) 8(26.7%) Vegetative 11(36.7%) 20(66.7%) 10(33.3%) 12(40.0%) 8(26.7%) Total 30(100.0%) 30(100%) 30(100%) 30(100%) 30(100%) Fig. 3. Plant number of tetrasporophyte and gametophyte in Chondrus ocellatus populations sampled on East and West coasts in between July and August 2013 (n=30 plants). Fig. 2. Average plant length (A) and weight (B) of Chondrus ocellatus sampled East and West coasts in between July and August 2013 (mean±sd, n=30 plants). 나, 서해안에서는학암포와만리포에서 36.7 % 와 46.7 % 로서해안에서높게나타났다. 과포자체는삼척에서 12개체 (40.0 %) 로최대였으며영덕에서가장적게관찰되었다. 생식기관을확인할수없는영양체를 acetalresorcinol test로사분포자체와배우체로구분하면, 동해안진두발개체군은배우체가 % 로서사분포자체 ( %) 에비해우점하였다. 하지만, 서해안의진두발개체군에서사분포자체비율 ( 만리포 70.0 %, 학암포 53.3 %) 은배우체비율 ( 만리포 30.0 %, 학암포 46.7 %) 에비해높게나타났다 (Fig. 3). 질병현황진두발의여름철개체군별 50개체중에서건강한조체는 11 21개체 ( %) 였으며, 질병에감염된개체는 29 39개체 ( %) 로서높게나타났다 (Fig. 4, Table 2). 진두발개체군에서관찰되는질병은검은반점 (black spot), 녹색반점 (green spot), 녹반증 (green colour), 백색증 (white colour) 로서연구정점에따라다르게나타났으며, 서해안만리포에서채집개체의 56.0 % 가검은반점을나타냈으며, 학암포개체의 32.0 % 는검은반점과녹반증을가지고있었다. 동해안의삼척, 영덕, 포항개체군에서질병조체의대부분이검은반점을가지는것으로확인되었다 (Table 2).

한국산진두발, Chondrus ocellatus 의생장과질병에대한연구 269 Fig. 4. Diseases of Chondrus ocellatus: A, Black spot; B, Green spot; C, White colour; D, Green colour. Scale bars represent 2 cm. Table 2.

Infected plants were classified into six groups State East Sea West Sea Samcheok Youngduk Pohang Hakampo Manripo Healthy plants 19(38.0%) 21(42.0%) 11(22.0%) 16(32.0%) 11(22.0%) Infected plants 31(62.

5 한국산진두발, Chondrus ocellatus 의생장과질병에대한연구 269 Fig. 4. Diseases of Chondrus ocellatus: A, Black spot; B, Green spot; C, White colour; D, Green colour. Scale bars represent 2 cm. Table 2. The number and percentage (%) of healthy and infected plants in Chondrus ocellatus populations collected on the rocky shores of East and West coasts, Korea. Infected plants were classified into six groups State East Sea West Sea Samcheok Youngduk Pohang Hakampo Manripo Healthy plants 19(38.0%) 21(42.0%) 11(22.0%) 16(32.0%) 11(22.0%) Infected plants 31(62.0%) 29(58.0%) 39(78.0%) 34(68.0%) 39(78.0%) Black spot(bs) 10(20.0%) 17(66.7%) 16(32.0%) 7(40.0%) 28(56.0%) Green spot(gs) 3(20.00%) - 14(28.0%) 4(8.0%) 2(4.0%) Green colour(gc) 8(16.0%) 11(22.0%) - 2(4.0%) 6(12.0%) White colour(wc) 1(2.0%) 1(2.0%) - 5(10.0%) 3(6.0%) BS+GS 5(10.0%) - 9(18.0%) - - BS+GC 4(8.0%) (32.0%) - Total 50(100.0%) 50(100.0%) 50(100.0%) 50(100.0%) 50(100.0%) -, no plant Fig. 5. Chondrus ocellatus with endophytic alga, Ulvella sp. (A, B) and epiphytic alga, Polysiphonia sp. (C-E).

6 270 이순정 박명애 아벨 박서경 김회경 김영식 최한길 진두발에서내생조류에감염되면엽체가녹색으로변화하였으며, 녹색으로탈색된개체에서내생조류로판단되는사상형녹조류 Ulvella sp. 가관찰되었다 (Fig. 5). 내생조류의숙주에대한감염과정과진두발의생존과생장에미치는영향은좀더심도깊은연구가있어야할것으로사료된다. 진두발에서착생하여서식하는해조류는붉은실류 (Polysiphonia sp.), 분홍잎류 (Acorsorium sp.), 잇바디가지류 (Herpochondria sp.), 애기가시덤불 (Caulacanthus okamurae), 나룻말류 (Acrochaetium sp.), 초록실 (Ulothrix flacca) 이었으며규조류도관찰되었다. 고찰한국연안에서식하는진두발 (C. ocellatus) 의길이와습중량은개체군에따라다르게나타났다. 동해안의삼척, 영덕, 포항의진두발개체군이서해안의만리포와학암포개체군에비해생장이좋았다. 진두발은카라기난을생산하는상업종인주름진두발 (C. crispus) 과같은속 (genus) 에속하는종으로서이들의길이생장과무게는단위면적당생산량과관련되므로카라기난산업에매우중요한정보이다. 포항의진두발은최대길이 cm까지, 영덕에서채집한개체는 g까지생장하므로향후진두발양식의모조를확보할때이곳의모조를사용하는것이생산량을높이는방법이다. 주름진두발의양식에서생산량을증대를위한최적생장조건탐색을위한배양연구 (Simpson and Shacklock, 1979) 가수행되었으며, Guiry (1981) 는다양한계통 (strain) 의생장을비교하여빠른생장을보이는암배우체 T4 를확보하였다. 한국산진두발은영덕개체군을제외하고동해안의삼척과포항, 그리고서해안의학암포와만리포에서 60% 이상의개체가생식기관을가지고있어여름철이성숙시기로사료된다. Kim et al. (2006) 은부산청사포의진두발개체군의성숙에대한 1년간의월 별연구에서 8월에 64 % 의엽체가성숙한것으로기록되어본연구와유사하였다. 영양체를 acetal-resorcinol test를실시한결과, 동해안의 3개진두발개체군은배우체와사분포자체비율이 1.7:1로나타났으나, 서해안의경우에는학암포에서 1:1.1, 만리포에서 1:2.3 로서동해안에서는배우체가, 서해안에서는사분포자체가우점하는것으로나타났다. 주름진두발 (C. crispus) 의배우체는상업적으로중요한젤리처럼굳어지는 κ-카라기난을포함하지만, 사분포자체는점성을가지며굳지않는 λ-카라기난을세포벽물질로가지고있어서생활사단계를파악하는것이중요하다. 동해안의경우에는상업적으로중요한배우체가서해안에비해높게나타났는데, 캐나다의 Prince Edward 섬에서는배우체가 % 정도로사분포자체에비해약간높게나타났다 (Lazo et al., 1989). Craigie and Pringle (1978) 은캐나다의동일한섬의다른지역에서배우체가 % 를차지한다고하였으며, 캐나다의 Nova Scotia의 Tor Bay에서 1994년에배우체와사분포자체비율은 3:1이라고하였으며 (Scrosati et al., 1994), 2011년에는지역에따라 1:1에서 9:1까지배우체가우점한다고하였다 (Garbary et al., 2011). 하지만, Lindgren and Åberg (1996) 는스웨덴의해양과기수지역의진두발개체군은사분포자체와배우체의비율이 1:1이라고하였다. 따라서, 진두발의배우체와사분포자체비율은해역과생활사에따라다르게나타나지만, 야외개체군에서배우체가사분포자체에비해우점하였다. 이러한이유는사분포자체가배우체에비해서내생해조류와병원체감염이쉽고섭식압이높기때문이라고하였다 (Bouarab et al., 2001; McLachlan et al., 2011). 하지만환경적스트레스지역, 즉기질 (substatum) 이불안정한해역에서는사분포자체와배우체비율이 1:1로유사하다고하였다 (Lazo et al., 1989). 본연구에서서해안태안반도의학암포와만리포에서배우체와사분포자체의비율이 1:1.1과 1:2.3로동해안의 1.7:1

7 한국산진두발, Chondrus ocellatus 의생장과질병에대한연구 271 과다르게나타나는원인을말하는것은많은연구가필요하겠지만, 서해안은수심이얕고갯벌이많아서퇴적물을포함한부유물질이많아동해안에비해서환경이안정되지않았기때문이아닌가사료된다. 우리나라연안에서식하는진두발의개체군에서현재까지질병에관한기록이없었으나, 본연구에서검은반점, 녹색반점, 녹반증, 백반증이엽체에서관찰되었으며, 내생조류 (endophytic algae) 와다양한착생조류가서식하고있는것으로밝혀졌다. 여름철의진두발은건강한조체 ( %) 에비해질병을가진개체는 % 로서높았다. 주름진두발에서 3종의내생녹조류가관찰되어분리하였으며 (Bown et al., 2003), Bouarab et al. (2001) 은 Chondrus crispus-acrochaete operculata 를숙주- 병원체 (host-pathogen) 의관계로언급하였고사분포자체가배우체에비해쉽게감염된다고하였다. 본연구에서관찰된사상형녹조류인 Acrochaete 속 (genus) 은현재 Ulvella 속의동종이명 (synonym) 으로여겨지고있으므로 (Guiry and Guiry, 2013), 진두발에서확인된종은 Ulvella sp. 로서유럽과캐나다에서발견되는병원체인내생조류가한국산진두발에서관찰되었으므로종동정, 감염경로및배우체가사분포자체에비해감염에대한저항성이있는지등에관한자세한연구가필요하다. 마지막으로진두발에서다양한착생해조류와규조류가관찰되었는데이들에대한계절적종다양성과풍도에대한연구가요구된다. 요약동해안의 3개정점 ( 삼척, 영덕, 포항 ) 과서해안의 2개정점 ( 학암포, 만리포 ) 에서여름철 (7, 8월 ) 에진두발을채집하여생장, 성숙, 배우체와사분포자체비율및질병에대한연구를실시하였다. 진두발의체장은개체군별로평균 cm였으며만리포에서최소였고포항에서최대로서동해안개체가서해 안개체에비해큰것으로확인되었다. 개체가서해안진두발에비해큰것을확인하였다. 진두발개체군의영양체비율은 % 였으며, 영양체를 resorcinol 방법으로구분한후배우체 : 사분포자체비율은동해안에서 1.7:1로배우체가우점하였고서해안에서는학암포에서 1:1.1, 만리포에서 1:2.3로서포자체가번무하였다. 본연구에서한국산진두발은다양한질병 ( 백색증과녹반증등 ), 내생조류와착생조류를가지고있었으며, 여름철의진두발은건강한조체는 % 로서많은개체가질병을가지고있었다. 진두발은현재까지동정되지않은사상형녹조류인내생하는 Ulvella sp. 를가지고있었다. 이러한내생조류는유럽과캐나다의주름진두발에서관찰되는감염성병원체로서보다많은관심과연구가요구된다. 감사의글본연구는국립수산과학원의지원 (RP-2013-AQ-217) 으로수행되었습니다. 참고문헌 Andrews, J.H.: The pathology of marine algae. Biol. Rev., 51: , Arévalo, R., Pinedo, S. and Ballesteros E.: Changes in the composition and structure of Mediterranean rocky-shore communities following a gradient of nutrient enrichment: Descriptive study and test of proposed methods to assess water quality regarding macroalgae. Mar. Poll. Bull., 55: , Amano, H., Suginaga, R., Arashim, K. and Noda, H.: Immunological detection of the fungal parasite,

8 272 이순정 박명애 아벨 박서경 김회경 김영식 최한길 Pythium sp.-the causative organism of red rot disease in Porphyra yezoensis. J. Appl. Phycol., 7: 53-58, Bates, S.S. and Craigie, J.S.: Fluorescence induction in the macroalgae Chondrus crispus (Rhodophyceae)and Ulva sp. (Chlorophyceae), Mar. Biol., 98: , Bown, P., Plumb, J., Sanchez-Baracaldo, P., Hayes, P.K. and Brodie, J.: Sequence heterogeneity of green (Chlorophyta) endophytic algae associated with a population of Chondrus crispus (Gigartinaceae, Rhodophyta). Eur. J. Phycol., 38: , Bouarab, K., Potin P., Weinberger F., Correa J. and Kloareg, B.: The Chondrus crispus-acrochaete operculata host-pathogen association, a novel model in glycobiology and applied phycopathology. J. Appl. Phycol., 13: , Braud, J.P. and Delepine, R.: Growth response of Chondrus crispus (Rhodophyta Gigartinales) to light and temperature in laboratory and outdoor tanks culture, pp In T. Levring (ed.) Proc. l0th Int. Seaweed Symp., Goteborg. W. de Gruyter, Berlin-New York, Brodie, J., Guiry, M. D. and Masuda, M.: Life history, morphology and crossability of Chondrus ocellatus forma ocellatus and C. ocellatus forma crispoides (Gigartinales, Rhodophyta) from the north-western Pacific, Eur. J. Phycol., 28: , Butardo, V.M., Ganzon-Fortes, E.T., Silvertre, VA., Bacano-Maningas, M.B.B., Montano, M.N.E. and Lluisma, A.O.: Isolation and classification of culturable bacteria associated with ice-ice disease in Kappaphycus and Eucheuma (Rhodophyta, Solieriaceae). Philipp. Schient., 40: , Choi, Y. and Kim, S.K.: Anticoagulant properties of carrageenans from Chondrus ocellatus. Proceedings of 1999 Spring Joint Meeting of the Korean Societies on Fisheries Science, pp , Chopin, T., Bodeau-Bellion, C., Floch, J.Y., Guittet, E. and Lallemand, J.Y.: Seasonal study of carrageenan structures from female gametophytes of Chondrus crispus Stackhouse (Rhodophyta). Hydrobiologia, 151/152: Correa, J.A. and Flores, V.: White rot disease in Gracilaria chilensis. J. Appl. Phycol., 7: , Craigie, J.S. and Correa, J.A.: Etiology of infectious diseases in cultivated Chondrus crispus (Gigartinales, Rhodophyta). Hydrobiologia, 326/327: , Cragie, J.S. and Leigh, C. Carrageenans and agars.: In Handbook of Phycological Methods: Physiological and Biochemical Methods, pp , Eds., Hellebust, J.A. and Craigie, J.S., Cambridge University Press, Cambridge, Craigie, J. S. and Pringle, J. D.: Spatial distribution of tetra-sporophytes and gametophytes in four maritime populations of Chondrus crispus. Can. J. Bot., 56: , Gall, Y.L., Braud, J. and Kloareg, B.: Protoplast production in Chondrus crispus gametophytes (Gigartinales, Rhodophyta). Plant Cell Reports, 8: , Garbary, D.J., Tompkins, E., White, K., Corey, P. and Kim, J.K.: Temporal and spatial variation in the distribution of life history phases of

9 한국산진두발, Chondrus ocellatus 의생장과질병에대한연구 273 Chondrus crispus (Gigartinales, Rhodophyta). Algae, 26: 61-71, Guiry, M.D.: Chondrus crispus Stackhouse T4 is a male clone (Rhodophyta). Phycologia, 20: , Guiry, M.D. and Guiry, G.M.: Algae Base. World-wide electronic publication, National University of Ireland, Galway. algaebase.org: Hattori, K., Kamohara, S., Harada, Y. and Kakinuma, M.: Detection of zoospores of the red rot disease fungus Pythium porphyrae by PCR in seawater from Porphyra cultivation farms at Aichi prefecture [Japan] in 2004 and Aqu. Sci., 55: , Ji, Y. and Guo, J.: The effect of temperature on the growth and development of Chondrus ocellatus. J. Dalian Fish. Coll. Dalian Shuichan Xueyuan Xuebao, 7: 32-37, Kang, J.W.: Illustrated Encyclopedia of Fauna & Flora of Korea Vol. 8. Marine Algae. Sam Hwa Press, Seoul, pp.465, Kang, J.W. and Koh, N.P.: Seaweed Cultivation. Tae Hwa Press, Pusan, pp. 294, Kim, Y.S., Choi, H.G. and Nam, K.W.: Phenology of Chondrus ocellatus in Cheongsapo near Busan Korea. J. Appl. Phycol., 18: , Largo, D.B.: Recent developments in seaweed diseases. In Proceedings of the National Seaweed Planning Workshop, pp , Eds., Hurtado, A.Q., Guanzon, N.G., de Castro-Mallare, T.R. and Luhan, M.R.J., SEAFDEC Aquaculture Department, Tigbauan, Iloilo, Largo, D.B., Fukami, K., Nishijima, T. and Ohno, M.: Laboratory-induced development of the ice-ice disease of the farmed red algae Kappaphycus alvarezii and Eucheuma denticulatum (Solieriaceae, Gigartinales, Rhodophyta). J. Appl. Phycol., 7: , Largo, D.B., Fukami, K., Adachi, M. and Nishijima, T.: Immunofluorescent detection of ice-ice disease-promoting bacterial strain Vibrio sp. P11 of the farmed macroalga, Kappaphycus alvarezii (Gigartinales, Rhodophyta). J. Mar. Biotechnol., 6: , Lazo, M.L., Greenwell, M. and McLachlan, J.: Population structure of Chondrus crispus Stackhouse (Gigartinaceae, Rhodophyta) along the coast of Prince Edward Island, Canada: distribution of gametophytic and sporophytic fronds. J. Exp. Mar. Biol. Ecol., 126: 45-58, Lee, Y.P. and Kang, S.Y.: A Catalogue of the Seaweeds in Korea. pp , Cheju National University Press, Cheju, Lindgren, A. and Ǻberg, P.: Proportion of life cycle stages of Chondrus crispus and its population structure: A comparison between a marine and an estuarine environment. Bot. Mar., 39: , Li, X., Jiang, Q., Lu, J. and Tao, W.: A description of Chondrus ocellatus Holmes and its variation in Bay of Liadong Peninsula. J. Dalian Fish. Coll. Dalian Shuichan Xueyuan Xuebao, 9: 21-25, Lüning, K.: Meeresbotank: Verbreitung, Ökophysiologie und Nutzung der marinen Makroalgen. Georg pp. 375, Thieme Verlag, Stuttgart., 1985.

10 274 이순정 박명애 아벨 박서경 김회경 김영식 최한길 McCandless, E. L., Craigie, J.S., Walter, J.A.: Carrageenans in the gametophytic and sporophytic stages of Chondrus crispus. Planta, 112: , McLachlan, J.L. Blanchard, W., Field, C. and Lewis, N.I.: Gametophyte life-history dominance of Chondrus crispus (Gigartinaceae, Rhodophyta) along the Atlantic coast of Nova Scotia, Canada. Algae, 26: 51-60, Narita, M., Sawabe, T., Gacesa, P., and Ezura, Y.: Rapid PCR detection of Pseudoalteromonas elyakovii, the causative bacterium of Laminaria spot-wound disease in Japan. Proceedings of the Int. Seaweed Symposium 17: , Orfanidis, S., Panayotidis, P. and Stamatis, N.: Ecological evaluation of transitional and coastal and water; A marine benthic macrophytes-based model. Medit. Mar. Sci., 2: 45-65, Park, C.S., Kakinuma, M. and Amano, H.: Detection of the red rot disease fungi Pythium spp. by polymerase chain reaction. Fish. Sci., 67: , 2001a. Park, C.S., Kakinuma, M., Amano, H.:Detection and quantitative analysis of zoospores of Pythium porphyrae, causative organism of red rot disease in Porphyra, by competitive PCR. J. Appl. Phycol., 13: , 2001b. Park, C.S.: Rapid detection of Pythium porphyrae in commercial samples of dried Porphyra yezoensis sheets by polymerase chain reaction. J. Appl. Phycol., 18: , Park, S.W., Lee, J.H. and Kim, Y.S.: Diseases of Seaweed. Bioscience Press, pp. 132, Scrosati, R., Garbary, D.J. and McLachlan, J.: Reproductive ecology of Chondrus crispus (Rhodophyta, Gigartinales) from Nova Scotia, Canada. Bot. Mar., 37: , Simpson, F.J. and Shacklock, P.F.: The cultivation of Chondrus crispus. Effect of temperature on growth and carrageenan production. Bot. Mar., 22: , Taylor, A.R.A. and Chen, L.C.M.: Chondrus Stackhouse, In Biology of Economic Algae, pp , Ed., Akatsuka, I., SPB Academic Publishing, Hague, Tribollet, A.D. and Vroom, P.S.: Temporal and spatial comparison of the relative abundance of macroalgae across the Mariana Archipelago between 2003 and Phycologia, 46: , Vasquez, J.A. and Guerra, N.: The use of seaweeds as bioindicators of natural and anthropogenic contaminants in northern Chile. Hydrobiologia, 326/327: , Zablackis, E., Vreeland, V., Doboszeski, B. and Laetsch, W.M.: Differential localization of carrageenan gelling sequences in Kappaphycus alvarezii var. tambalang (Rhodophyta) with FITC-conjugated carrageenan oligosaccharides. J. Phycol., 27: , Manuscript Received : October 21, 2013 Revised : November 14, 2013 Accepted : November 12, 2013

03이경미(237~248)ok

03이경미(237~248)ok The recent (2001-2010) changes on temperature and precipitation related to normals (1971-2000) in Korea* Kyoungmi Lee** Hee-Jeong Baek*** ChunHo Cho**** Won-Tae Kwon*****. 61 (1971~2000) 10 (2001~2010).