식물병연구 Note Open Access Res. Plant Dis. 22(4): (2016) 토마토품종의토마토황화잎말림바이러스병에대한저항성평가 Resistance Evaluati

식물병연구 Note Open Access Res. Plant Dis. 22(4): 297-302 (2016) https://doi.org/10.5423/rpd.2016.22.4.297 토마토품종의토마토황화잎말림바이러스병에대한저항성평가 Resistance Evaluation of Commmercial Tomato Cultivars against Tomato yellow leaf curl virus 고숙주 1 * ㆍ김효정 1 ㆍ이진희 1 ㆍ마경철 1 ㆍ최덕수 1 ㆍ박영훈 2 ㆍ최승국 3 ㆍ김미경 4 ㆍ최홍수 4 1 전라남도농업기술원친환경농업연구소, 2 부산대학교원예생명과학과, 3 농촌진흥청국립원예특작과학원원예특작환경과, 4 농촌진흥청국립농업과학원작물보호과 *Corresponding author Tel: +82-61-330-2512 Fax: +82-61-336-4076 E-mail: kosj@korea.kr Sug-Ju Ko 1 *, Hyo-Jeong Kim 1, Jin-Hee Lee 1, Kyung-Cheol Ma 1, Duck-Soo Choi 1, Young-Hoon Park 2, Seung-Kook Choi 3, Mi Kyeong Kim 4, and Hong-Soo Choi 4 1 Environment-friendly Agricultural Research Institute, Jeonnam Agricultural Research & Extension Services, Naju 58213, Korea 2 Department of Horticultural Bioscience, Pusan National University, Miryang 50463, Korea 3 Horticultural and Herbal Crop Environment Division, National Institute of Horticultural and Herbal Science, Rural Development Administration, Wanju 55365, Korea 4 Crop Protection Division, National Institute of Agricultural Sciences, Rural Development Administration, Wanju 55365, Korea Received September 10, 2016 Revised November 10, 2016 Accepted November 11, 2016 Tomato yellow leaf curl virus (TYLCV) is a viral disease causing severe economic losses on tomato. Practical prevention of the TYLCV disease is to control tabacco whitefly (Bemisia tabaci) or to cultivate TYLCV-resistant tomato cultivars, because no agrochemical products are available to control TYLCV. In this study, TYLCV resistance of the commercial tomato cultivars were evaluated using the DNA markers tightly linked to TYLCV resistance genes Ty-1 and Ty-3 and infection with the TYLCV clones mediated by Agrobacterium. In marker genotyping, resistance alleles were detected from 4 oval type tomato cultivars (Titichal, TY tinny, TY saengsaeng II, TY sense Q). Four cheery type cultiavrs (TY endorphin, TY smartsama, Tiara TY, Olleh TY) and 6 round type cultivars (TY kingdom, TY ace, TY homerun, TY altorang, Dotaerang TY winner, Styx TY). The seedling bioassay indicated that tomato cultivars of the oval type and cherry type showed consistancy in marker genotype and phenotype while slight disease symptom was observed from some round type cultivras (TY ace, TY homerun, Styx TY) with resistance marker genotype. For fruit yields, TY tinny was greater than its control cultivar Titichal in oval types, TY smartsama was greater than its control Smile in cherry type, and TY ace and TY kingdom were greater than their control Dabok. These cutliavrs can be a good choice for high-yielding TYLCVresistant tomato cultivars. Keywords: Cultivar, Resistance, Tobacco whitefly, Tomato, Tomato yellow leaf curl virus 토마토황화잎말림바이러스 (Tomato yellow leaf curl virus, TYLCV) 는 Geminiviridae 과, Begomovirus 속이며미국, 남미, 유 Research in Plant Disease pissn 1598-2262, eissn 2233-9191 www.online-rpd.org 럽, 동남아시아, 일본, 중국등전세계토마토재배지에피해를초래하고있다 (Czosnek와 Laterrot, 1997; Hanssen 등, 2010; Navas-Castillo 등, 2011). 국내에서는경상남도통영의토마토농가에서 2008년 5월처음발생되었고 (Ji 등, 2008; Kwak 등, 2008), 2010년에는경남통영등 16개시군, 경북 10개, 충남 4 The Korean Society of Plant Pathology cc This is an open access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/ licenses/by-nc/4.0/), which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited. 297

298 Research in Plant Disease Vol. 22 No. 4 개, 충북 2개, 전남 13개, 전북 11개, 제주도 2개시군등전국적으로급속히확산되었다 (Kim 등, 2011). TYLCV는 1964년이스라엘에서처음보고된후분리지역, 병징차이등에따라 10여종의계통이확인되었다 (Abhary 등, 2007; Cohen과 Harpaz, 1964; Fauquet 등, 2003). TYLCV에감염되면황화, 잎말림및위축등의증상을보이고, 담배가루이가 TYLCV를직접매개하며, 즙액, 종자및접촉전염은하지않는다고하였다 (Czosnek, 1999; Diener와 Raymer, 1971). 담배가루이는고온기에세대가짧아지고, 발생횟수가많아져살충제에대한약제저항성이발달하여방제가잘되지않으므로 (Devine 와 Denholm, 1998; Lee 등, 2012; Nauen 등, 2002) 방충망을이용한시설내유입차단방법과트랩식물을이용한방제법이보고되었다 (Choi 등, 2016; Chung 등, 2013). 하지만방충망이용방법은고온기에온도관리의한계가있어근본적인바이러스방제대책이되지않는다. 품종을선택할때매개충의주발생시기를고려하여촉성재배는품종의저항성여부에제한을두지않고, 담배가루이밀도가증가하는 7월이후정식시에는저항성품종재배를권장하였다 (Ko 등, 2014). 지금까지알려진 TYLCV 저항성유전자는 Ty-1, Ty-2, Ty-3, Ty- 3a, Ty-4, Ty-5이며, 최근유전자지도분석에따르면 Ty-1과 Ty-3 이매우근접하게위치하여동일한유전자좌의대립유전자일가능성이있다고하였다 (Anbinder 등, 2009; Garcia 등, 2007; Hoogstraten과 Braun, 2005; Ji와 Scott, 2006; Ji 등, 2007, 2009; Verlaan 등, 2011). TYLCV 저항성품종개발을위한유전자마커 Ty-1, Ty-2, Ty-3, Ty-3a가육종에이용되고있다 (Hoogstraten과 Braun, 2005; Ji 등, 2007). 현재시중에유통되고있는 TYLCV 내병성품종들이농가에서재배되고있으나이에대한연구로는 Kim 등 (2013) 이발표한유럽계 34품종에대한저항성정도와수량성등에대한보고뿐이다. 본연구는전남지역에서재배되고있는토마토품종의 TYLCV 저항성유전자보유여부와 TYLCV 를접종하였을때표현형을조사하고자하였으며, 내병성품종들의원예형질특성을조사하여영농현장에서품종선택의기초자료로제공하고자하였다. TYLCV 저항성검정. 토마토품종은일반 ( 완숙 ) 형, 방울형, 대추형 3가지형태로구분하여총 14개내병성품종과 8 개감수성품종을대상으로실시하였다. 일반 ( 완숙 ) 형은과중이 150 250 g 크기의원형이고, 방울형은 15 20 g 크기의작은원형이며, 대추형은 15 20 g 크기의타원형이다. TYLCV 접종은국립원예특작과학원바이러스실에서 TYLCV 감염성클론을분양받아 Choi 등 (2013) 의방법에따라실시하였 다. 형질전환된 Agrobacterium tumefaciens를 Kanamycin 등 3 종의항생제를첨가한 Luria-Bertani (LB) 배지에서 28 o C, 24시간진탕배양하였다. 배양액 10 μl를 10 ml의 LB 배지에첨가한후 28 o C, 8시간진탕배양하였다. 배양액을 4,500 rpm에서 10분간원심분리후회수하여 OD 600 0.7 0.8 농도로현탁하였다. 현탁된균에 10 mm MgCl 2, 200 μm acetosyringone, 10 mm MES를넣어서상온에서 2시간가량둔후에 1 ml 주사기를사용하여아그로주입법 (Choi 등, 2013) 으로접종하였다. 토마토종자를파종하여 2 3엽기에개체당떡잎 2개의잎뒷면에균 1 ml를주입하여접종하였다. 접종한식물체는 25 o C±5 o C로유지되는온실에서 30일동안재배하면서병징을조사하였다. 바이러스접종은품종당 10주씩 3반복으로실시하였다. 바이러스조사는 Choi 등 (2013) 의방법을변형하여증상없음 (0), 약한증상 (1), 황화증상 (2), 위축과황화잎말림 (3) 의 4단계로구분하고다음과같이발병도로환산하였다. 발병도 (%)=[ ( 발병수 발병계수 )/3 개체수 ] 100. TY- LCV 내병성유전자를확인하기위한검정은 Hwang 등 (2012) 과 Kim 등 (2013) 의방법에따라수행되었다. 품종별로유묘기식물체의본엽에서 total genomic DNA를추출한후 TYLCV 저항성유전자 Ty-1과 Ty-3과연관된분자마커 (Kim 등, 2013) 를이용하여확인한결과 (Table 1), 대추형은티티찰, TY 티니, TY 센스큐 3품종에서 Ty-1과 Ty-3을확인하였고, TY 생생 II에서는 Ty-1만확인되었다. 방울형은 TY 엔도르핀, TY 스마프사마, 티아라 TY, 올레 TY 4종에서 Ty-1과 Ty-3 유전자를확인하였다. 완숙형은 TY 에이스, TY 홈런, TY 알토랑, 도테랑 TY 위너, 스틱스 TY 등 5종에서 Ty-1과 Ty-3 유전자를확인하였으나 TY 킹덤은 Ty-1 유전자만확인되었다. 따라서 Ty계통에서는모두 Ty 유전자가이형접합으로존재함이확인되었으나조사된 8개감수성품종에서는모두저항성유전자가확인되지않았다 (Table 1). 생물검정에서는대추형티티찰, TY 티니, TY 생생 II, TY 센스큐등저항성품종 4종에서병징이나타나지않았으나감수성품종미니찰에서 12.2%, 베타티니에서 23.3% 의발병도를나타냈다. 방울형은 TY 엔도르핀, TY 스마프사마, 티아라 TY, 올레 TY 등저항성품종 4종에서병징이나타나지않았으나감수성품종인스마일 27.8%, 티아라 31.1%, 유니콘 41.1% 의발병도를보였다. 완숙형은저항성품종중 TY 킹덤과 TY 알토랑에서는병징을보이지않았으나 TY 에이스는 33.3%, TY 홈런 10.0%, 도테랑 TY 위너 2.2%, 스틱스 TY 7.8% 의발병도를나타냈다. 감수성품종에서는마스카라맵시 77.8%, 호용 77.8%, 다복 90.0% 의발병도를나타냈다. 완숙형 4종의저항성품종에서병징이발현된원인은정확히알수없으나저항성유전자가존재하더라도병원균의밀도가

Research in Plant Disease Vol. 22 No. 4 299 Table 1. Results of bioassay and molecular marker evaluation of tomato cultivars for the resistance to Tomato yellow leaf curl virus Type Varieties Company Ty-1 Resistance gene* Ty-3 Disease severity (%) Oval Titichal Nongwoo H H 0 a TY Tinny PPS H H 0 a TY Saengsaeng II Namnong H S 0 a TY Sense Q Nongwoo H H 0 a Minichal Nongwoo S S 12.2 b Betatini PPS S S 23.3 c Cherry TY Endorphin Bunong H H 0 a TY Smartsama Coregon H H 0 a Tiara TY PPS H H 0 a Olleh TY Dongbufarm H H 0 a Smile Namnong S S 27.8 b Tiara PPS S S 31.1 bc Unicorn Dongbufarm S S 41.1 c Common TY Kingdom Bunong H S 0 a TY Ace Namnong H H 33.3 b TY Homerun Namnong H H 10.0 a TY Altorang Nongwoo H H 0 a Dotaerang TY Winner Nongwoo H H 2.2 a Styx TY Dongbufarm H H 7.8 a Mascaramaepssi Bunong S S 77.8 c Hoyong Sakada S S 77.8 c Dabok Namnong S S 90.0 c *H, heterozygous marker genotype indicating dominant effect of resistance; S, homozygous maker genotype for susceptible allele. Disease severity from 0 to 3. 0, healthy looking; 1, mild symptom; 2, yellowing symptom; 3, stunt with severe symptom. Disease severity (%)=[ (number of diseased plant disease index)/3 number of plant] 100. Means followed by a common letter are not significantly different at the 5% by Duncan s multiple range test. 높거나병원성이증가되는환경에서는병징이나타난다는보고도있다 (Kim 등, 2013). 또한사용된마커가저항성유전자기반의마커가아닌유전자연관마커임을고려할때이들품종에서는마커-유전자간재조합이일어났을가능성이있다. 하지만감수성품종에비해서는발병도가매우낮게나타난사실을볼때전자의가능성이보다높을것으로판단된다. 품종별생육및수량성검정. 본시험은대추형품종인 TY 생생 II, TY 티니, TY 센스큐, 티티찰 4종, 방울형인 TY 스마프사마, 티아라 TY 2종, 완숙형인도테랑 TY 위너, TY 킹덤, TY 홈런, TY 에이스, TY 알토랑 5종을공시하여전남나주시소재전남농업기술원비닐하우스에서수행하였다. 정 식은 2014년 4월 17일에 1화방의 1번꽃이개화하기직전에 70 25 cm의재식거리로 2열로정식한후점적호스를설치하고흑색비닐로멀칭하였다. 토마토는 5화방까지재배하여적심하였고모든측지는제거하였다. 수분은나투벌 (Koppert, Monster, The Netherlands) 을이용하였으며, 재배지의토양은식양토로시비방법은밑거름 11.6-10.3-4.1-2000 (N-P-K-퇴비) kg/10 a 시용하였고, 추비는 8.8-8.1 (N-K) kg/10 a 3회에걸쳐분시하였다. 수확기간은 6월 9일부터 9월 5일까지이며, 과실의무게와상품수량을화방별로분리하여조사하였고수용성고형물함량 ( 당도 ) 은굴절당도계 (ATC-20E; Atago, Tokyo, Japan) 를사용하여측정하였다. 생육특성은초장, 절간장, 엽장, 엽폭을조사하였고, 완전임의배치 3반복으로하여반복당 20주를공시하였다.

300 Research in Plant Disease Vol. 22 No. 4 생육특성을살펴보면초장은 TY 생생 II 236 cm, 티티찰 203 cm, TY 티니 193 cm, TY 센스큐 183 cm 순이었고, 대비품종인미니찰은 176 cm로가장작았다. 방울형은티아라 TY 194 cm, TY 스마프사마 191 cm, 대비품종인유니콘 197 cm, 스마일 154 cm였다. 완숙형은 TY 킹덤 200 cm, TY 알토랑 193 cm, TY 홈런 177 cm, 도테랑 TY 위너 173 cm, TY 에이스 167 cm, 대비품종인마스카라맵시 197 cm, 다복 191 cm였다. 절간장도비슷한경향이었다 (Table 2). 과중은대추형은 TY 티니 17.6 g, 티티찰 16.6 g, TY 생생 II 16.0 g, TY 센스큐 15.8 g, 대비품종인미니찰 15.7 g 순이었으나유의차는없었다. 방울형은 TY 스마프사마 16.7 g, 티아라 TY 16.3 g, 대비품종인스마일 14.7 g, 유니콘 13.4 g이었으며, TY 스마프사마와티아라 TY가대비품종에비해큰것으로유의차가있었다. 완숙형은 TY 킹덤 224.1 g, TY 알토랑 189.3 g, TY 에이스 170.7 g, TY 홈런 159.4 g, 도테랑 TY 위너 136.6 g, 대비품종인다복 184.1 g, 마스카라맵시 159.2 g이었으며, TY 킹덤 224.1 g과 TY 알토랑은다복과비슷한대과종, TY 에이스와 TY 홈런은마스카라맵시와비슷한중과종, 도테랑 TY 위너는소과종이었다 (Table 3). 당도는대추형인 TY 생생 II 7.6 o Bx, TY 티니 7.2, 티티찰 7.2, TY 센스큐 7.1, 대비품종인미니찰 7.5로 TY 생생 II를제외하고모두낮았다. 방울형인티아라 TY 6.2, TY 스마프사마 5.7로 대비품종인스마일 7.0, 유니콘 6.8에비해모두낮았다. 완숙형인 TY 킹덤 4.4, TY 알토랑 4.4, TY 에이스 3.9, TY 홈런 3.6, 도테랑 TY 위너 4.8, 대비품종인다복 4.4, 마스카라맵시 4.8이었으며, 도테랑 TY 위너와 TY 킹덤은대비품종과유사하였으나 TY 에이스와 TY 홈런은대비품종에비해낮은편이었다 (Table 3). 수량은대추형인티티찰 3,656 kg, TY 티니 3,419 kg, TY 센스큐 3,124 kg, TY 생생 II 1,733 kg, 대비품종인미니찰 2,549 kg 으로티티찰, TY 티니, TY 센스큐는미니찰보다많았다. 방울형인 TY 스마프사마 2,880 kg, 티아라 TY 2,861 kg, 대비품종인스마일 1,439 kg, 유니콘 621 kg이었으며, TY 스마프사마와티아라 TY가대비품종에비해많았다. 완숙형인 TY 에이스 4,476 kg, TY 킹덤 4,196 kg, TY 홈런 3,863 kg, TY 알토랑 3,503 kg, 도테랑 TY 위너 2,577 kg, 대비품종인다복 3,972 kg, 마스카라맵시 2,106 kg이었으며, TY 에이스, TY 킹덤, TY 홈런은다복과비슷하였다 (Table 3). Ko 등 (2014) 이제시한경종적방제법을고려한다면 TYLCV 주발생시기에는저항성품종을재배하여야한다. 수량성을고려한다면대추형은 TY 티니, 방울형은 TY 스마프사마, 티아라 TY, 완숙형은 TY 에이스, TY 킹덤을우수한품종으로추천한다. Table 2. Growth characteristics of tomato cultivars evaluated for the resistance to Tomato yellow leaf curl virus (TYLCV) Type Varieties Plant height (cm) Internode length (mm) Stem diameter (mm) Position of first fruit set Leaf length (cm) Leaf range (cm) Oval TY Saengsaeng II 235.8 a* 73.2 a 13.0 b 9.3 a 42.4 ab 37.7 a TY Tinny 193.3 bc 52.7 c 17.5 ab 7.1 b 44.5 ab 37.5 a TY Sense Q 182.8 bc 50.4 c 16.6 ab 6.7 b 40.5 ab 36.4 a Titichal 203.1 b 60.4 b 17.8 a 6.2 b 45.5 a 38.3 a Minichal 175.7 c 50.2 c 18.2 a 6.9 b 39.1 b 38.3 a Cherry TY Smartsama 190.8 a 60.9 ab 14.5 a 6.1 a 35.5 b 33.7 b Tiara TY 193.6 a 65.2 ab 16.4 a 5.9 a 37.1 ab 30.8 b Smile 154.3 b 46.7 b 17.3 a 4.7 a 41.7 a 44.6 a Unicorn 196.6 a 61.9 ab 14.9 a 6.0 a 37.9 ab 34.5 b Common Dotaerang TY Winner 173.1 bc 75.0 bc 15.3 ab 6.9 a 39.9 a 42.1 ab TY Kingdom 200.2 a 84.6 ab 16.9 a 7.1 a 41.5 a 44.3 ab TY Homerun 177.7 bc 71.7 c 14.6 ab 8.2 a 37.2 a 37.5 b TY Ace 166.5 c 66.9 c 13.8 b 6.5 a 40.2 a 41.1 ab TY Altorang 193.1 ab 88.9 a 15.8 ab 7.4 a 43.0 a 48.9 a Dabok 190.6 ab 82.3 ab 14.5 ab 6.5 a 40.1 a 44.1 ab Mascaramaepssi 197.1 a 82.6 ab 14.2 b 7.8 a 39.3 a 44.7 ab *Means followed by a common letter are not significantly different at the 5% by Duncan s multiple range test.

Research in Plant Disease Vol. 22 No. 4 301 Table 3. Fruit quality of tomato cultivars evaluated for the resistance to to Tomato yellow leaf curl virus (TYLCV) Type Varieties Fruit weight (g) Sugar content (brix) Total yield (kg/10 a) Good yield (kg/10 a) Oval TY Saengsaeng II 16.0 a* 7.6 a 3,033 b 1,733 b TY Tinny 17.6 a 7.2 bc 4,482 a 3,419 ab TY Sense Q 15.8 a 7.1 c 4,240 ab 3,124 ab Titichal 16.6 a 7.2 bc 4,983 a 3,656 a Minichal 15.7 a 7.5 ab 3,897 ab 2,549 ab Cherry TY Smartsama 16.7 a 5.7 d 4,467 a 2,880 a Tiara TY 16.3 a 6.2 c 4,814 a 2,861 a Smile 14.7 ab 7.0 a 3,709 a 1,439 ab Unicorn 13.4 b 6.8 b 4,076 a 621 b Common Dotaerang TY Winner 136.6 c 4.8 a 5,393 a 2,577 a TY Kingdom 224.1 ab 4.4 ab 5,402 a 4,196 a TY Homerun 159.4 bc 3.6 c 7,428 a 3,863 a TY Ace 170.7 bc 3.9 bc 7,374 a 4,476 a TY Altorang 189.3 ab 4.4 a 5,621 a 3,503 a Dabok 184.1 b 4.4 a 6,905 a 3,972 a Mascaramaepssi 159.2 bc 4.8 a 4,789 a 2,106 a Trantsplanting: April 17, 2014; harvesting period: June 9, 2014 to September 5, 2014. *Means followed by a common letter are not significantly different at the 5% by Duncan s multiple range test. 요약 Conflicts of Interest 토마토황화잎말림바이러스 (TYLCV) 는토마토에서큰경제적손실을초래하는바이러스병이다. 이병은약제방제가되지않기때문에매개충인담배가루이를방제하거나저항성품종을재배해야한다. 본시험은시중에유통중인토마토품종에대해토마토황화잎말림바이러스에저항성을평가하였다. 토마토품종별로 TYLCV 저항성마커로유전자 Ty-1 과 Ty-3 분석을실시하였고, 아그로주입법으로생물검정을실시하였다. 대추형은티티찰, TY 티니, TY 생생 II, TY 센스큐등 4종, 방울형은 TY 엔도르핀, TY 스마프사마, 티아라 TY, 올레 TY 등 4종, 완숙형은 TY 킹덤, TY 에이스, TY 홈런, TY 알토랑, 도테랑 TY 위너, 스틱스 TY 등 6종에서저항성유전자를확인하였다. 유묘검정은대추형과방울형은모두유전자분석결과와일치하였으나, 완숙형은저항성품종에서도일부병징이발현되는경향이었다. 품종별수량성은대추형은티티찰대비 TY 티니가우수하였고, 방울형은스마일대비 TY 스마프사마, 티아라 TY이우수하였으며, 완숙형은다복대비 TY 에이스, TY 킹덤등이우수한품종이었다. No potential conflict of interest relevant to this article was reported. Acknowledgement This study was carried out with the support of Cooperative Research program for Agricultural Science & Technology Development (Project No. PJ01011307) Rural Development Administration, Republic of Korea. References Abhary, M., Patil, B. L. and Fauquet, C. M. 2007. Molecular biodiversity, taxonomy, and nomenclature of tomato yellow leaf curl-like viruses. In: Tomato Yellow Leaf Curl Virus Disease, ed. by H. Czosnek, pp. 85-118. Springer, Dordrecht, Netherlands. Anbinder, I., Reuveni, M., Azari, R., Paran, I., Nahon, S., Shlomo, H., Chen, L., Lapidot, M. and Levin, I. 2009. Molecular dissection of Tomato leaf curl virus resistance in tomato line TY172 derived from Solanum peruvianum. Theor. Appl. Genet. 119: 519-530. Choi, S. G., Choi, H. S., Yang, E. Y., Cho, I. S., Cho, J. D. and Chung,

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