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Res. Plant Dis. 16(2) : 141-147 (2010) Research in Plant Disease The Korean Society of Plant Pathology ü q š t w ½ *Á«k 1,2 Áy 1 Á Á Á xá½x w w, 1» š x, 2 x» t» x Resistance to Phytophthora Blight of Commercial Pepper Cultivars in Korea Byung-Soo Kim*, Tae-Ryong Kwon 1,2, Ji-Eun Hwang 1, Jae-Moo Lee, Dong-Guen Park, Joon-Hyung Ahn and Hye-Yeon Kim Department of HorticulturalG Science, Kyungpook National University, Daegu 702-701, Korea 1 Youngyang Pepper Experiment Station, Gyeongsangbuk-do Agricultural Research and Extension Services, Yeongyang 764-803, Korea 2 Current address: Punggi Ginseng Experiment Station, Gyeongsangbuk-do Agricultural Research and Extension Services, Yeongju 750-871, Korea (Received on July 20, 2010; Accepted on July 25, 2010) Resistance to Phytophthora blight (Phytophthora capsici Leonian) of commercial cultivars of Capsicum pepper in Korea were evaluated from 2008 to 2010. In 2008 and 2009, the cultivars were tested for resistance to the Pc003 isolate collected in Youngyang, Gyeongbuk province. In 2010, the selected cultivars in the previous years were tested by inoculating with 3 isolates obtained from the pepper plants in Miryang in Gyeongnam province, Youngyang in Gyeongbuk province, and Goesan in Chungbuk province. A continuous variation in resistance from highly resistant to susceptible was observed among the commercial pepper cultivars. It was also noted that some cultivars, although their names were initiating with Yeokgang or PR meaning Phytophthora resistance, were very low in resistance or susceptible. When the resistant cultivars selected in 2008 and 2009 experiment were inoculated with the 3 isolates, all the commercial cultivars except a rootstock, Tantan, succumbed to the exceptionally virulent Pc005 (Miryang) isolate. Pc002 (Goesan) was a little more virulent than Pc003 (Youngyang). A few cultivars resistant to Pc003 (Youngyang) were severely infected by Pc002 (Goesan). Significant interaction in analysis of variance suggested the differential interactions between cultivars and pathogen isolates. Strategies to breed cultivars having high level of resistance to the highly variable pathogenic fungus, Phytophthora capsici, were discussed. Keywords : Capsicum annuum, Differential reaction, Phytophthora capsici, Resistance 2008 m š ƒ 4 8 8 ha, 6 ha w w 5 4 8 ha w. š ƒ 9,117, tš ƒ 4,048 ww 1 3 165 w 2 wš ( t, www.maf.go.kr). p,,, y š x. ù š w t m *Corresponding author Phone) +82-53-950-5729, Fax) +82-53-950-5722 Email) bskim@knu.ac.kr w vw š. š w», s w ù, ³ m. w t ƒ. š w w, (Bosland Lindsey, 1991; ½, 1986, 1988; ½, 1975; Kim, 2001; Kimble Grogan, 1960), w t w t w w (½, 1992; ½, 1996). ü š t z wš, w t ƒ

142 ½ Á«k Áy Á Á Á xá½x» w 2005 l. z w ü t q š w t w ƒ, ³ w t ³ (Virulence) y ƒ w j. 2008 l ü q ù q t w w sƒwš t 3 ³ w w t wš w. 2008 : q y t w. q t w sww 70 œ 2 18 q z 3 6 36 sp w z, q z 60 4 21 1.5 10 /ml ³ s xk 5 w. ƒ t 36 w. w Pc003 ³ w. PDA ³ ³ V8 (Agar 20 g, V8 Juice 200 ml, CaCO 3 3 g, Distilled Water 800 ml) e w z 24 o C xÿw 10 w s û x w. s xk s û x t s û xk 2 w z 7C 1 þ z o 3 g s û l s g w. s xk x» w 1.5 10 /ml 5 w. s xk Dispenser w ƒ 5ml w. 9 z ù w. s 1.8 m n PE v š w sp z w š ü Ÿ w 28 o C, 20 o C w. 2009 : 2008 t w. qt 55 œ 2 14 128 p q w. q z 30 3 17 32 p w z, q z 55 4 11 ³ s û xk w. ³ w x Pc003 ³ w. V8 ³ ³ j y 5 5 mm, ¾ 5 mm 5 ü V8 ³ w. ü y 5 ( ) üš û y t v r ³ ƒ w. w 24 o C 4-5 wì y t s x w. y t s û x w k s û xk w. s ûxk 2 w z s û 5 10 s û/ml 5 w. ƒ s ûxk 5ml w w. ³ wš 16 z w w.» 1=» ( ) ; 2=» ù š ; 3= w š, 4= 4 w. w 1= ; 2= 25% ; 3= 50% ; 4= 75% ; 5= 100% 5 w. 2010 : t 3 ³ w w. qt 36 œ 3 13 200 p q w q z 30 4 16 50 p w. 5 3 50 z t 3 š ³ w w w. ³ Pc002( ), Pc003( ), Pc005( ) 3 ³ 2009 y w t x s û w. 21 z w 2009» w. š 2008 : q y t w. 71 œ t ùkü Table 1, ƒ ùkù t Table 2 ù kù. w ƒ ¾ k û t 26 37.1% w. t 1-2 ƒ t l ƒ t ¾ w w ùkü. p, PR

ü q š t w 143 Table 1. Frequency of commercial hybrid cultivars by percentage of diseased plants 9 days after inoculation by drenching zoospore suspension of isolate Pc003 of Phytophthora capsici Category in percentage of diseased plants (PDP) Frequency of cultivars Percent frequency of cultivars PDP=0 26 37.1 0<PDP 20 8 11.4 20<PDP 50 5 7.1 PDP>50 31 44.3 Total 70 99.9 Table 2. Twenty six commercial cultivars ranked top with no diseased plants 9 days after inoculation by drenching zoospore suspension of isolate Pc003 of Phytophthora capsici Cultivar Cheonnyeonmannyeon, Daewanggeon, Dokyacheongcheong, Gangryeokdaetong, Gangryeokjosenggeon, Geumgochu, Geumnara, Giripbaksu, Ildangbaek, Ildeunggongshin, Ilsongjeong, Mansahyeongtong, Muhandojeon, Muhanjilju, PR-Daechon, PR- Galmuri, PR-Sansaeng, PR-Taepyeong, Shinhwachangjo, Taesan, HNX14460212, HNX14460242, N6501, N6510, N6519, N6538 w w ƒ t w ƒ 50% y 100% t. q š š t w ü ù w w w ù t m š ùkü. ¾ ü š t w sƒw t w q w» ù ƒƒ w t vw ƒ. w w w» w» x ü q w w œ w v w. š ƒ w ù ü w ù mw w. 2009 : 2008 t w. 2008 x w t,, w p š w w t t 55 œ w w w w ƒ w 27t t Table 3 ùkü.» ƒ 5t w, PR-,,, k. y,, œ, PR-,, PR-,» ùkùš. x Table 3. Resistance to Phytophthora blight of commercial cultivars of pepper ranked about upper half in resistance of 63 cultivars tested in 2009 Commercial No. Plants Disease index Cultivar inoculated Stem rot z Root rot y Muhanjilju 16 1.0 a x 1.0 a PR-Manse 16 1.0 a 1.2 ab Ilsongjeong 16 1.0 a 1.3 a-c Cheonnyeonmannyeon 16 1.0 a 1.4 a-d Taesan 16 1.0 a 1.5 a-e Yeokganghongjanggun 16 1.0 a 1.6 a-f Ilinja 16 1.0 a 1.8 b-f Ildeunggongshin 16 1.0 a 1.9 d-f PR Sadaecheonwang 16 1.0 a 2.8 g Josaenggeomul 16 1.0 a 3.0 g PR Sangsaeng 16 1.1 a 1.3 a-c Giripbaksu 16 1.1 a 1.6 a-f PR-Oraetta 16 1.1 a 2.0 ef Ildangbaek 16 1.1 a 2.6 g Seonguja 16 1.1 a 1.2 ab PR-Galmuri 16 1.1 a 2.1 f PR-Yeoljeong 16 1.2 ab 1.2 ab Mansahyeongtong 16 1.3 ab 1.5 a-e PR-Bultina 16 1.3 ab 2.8 g PR-hongdukkae 16 1.3 ab 2.9 g Shintobuli 16 1.4 b 1.4 a-d Gangryeokjosenggeon 16 1.8 c 1.8 c-f 6510 16 1.9 cd 1.3 a-c Gangryeokdaetong 16 1.9 cd 1.6 a-f PR Taepyeong 16 2.0 cd 1.5 a-e Daegeonwang 16 2.1 cd 1.3 a-d HNX14460242 16 2.1 cd 1.8 c-f Shinhwa (Control) 16 4.0 e 5.0 h x Mean separation within columns by Duncan's multiple range test, P 0.05. y 1 = No root rot observed; 2 = about 25% root rot; 3 = about 50% root rot ; 4 = about 75% root rot; 5 = complete root rot. z 1 = No disease symptoms observed; 2 = necrotic lesion at base of stem, yet still surviving; 3 = wilting; 4 = dead and dried. ³ Pc003( ) 1 w» w ³. w x w ùkù t ³ w, s k w w w. 2010 : t 3 ³ w w. 2008-2009 x ñ t sww

144 ½ Á«k Áy Á Á Á xá½x Table 4. Resistance of commercial cultivars of pepper selected for resistance in 2008-2009 to 3 isolates of Phytophthora capsici in 2010 test Cultivar No. Plants Pc002 (Goesan) Pc003 (Youngyang) Pc005 (Miryang) inoculated Stem Root Stem rot Root rot Stem rot z Root rot y Tantan 15 1.0 a 1.9 a-c 1.0 a 1.0 a 1.0 a x 3.9 a PR-Yeoljeong 15 1.0 a 1.8 ab 1.0 a 1.0 a 4.0 d 5.0 d Geumgochu 15 1.1 ab 2.1 a-d 1.0 a 1.0 a 3.9 d 4.9 cd Shintobuli 15 1.1 ab 2.5 b-e 1.0 a 1.0 a 4.0 d 5.0 d Ilsongjeong 15 1.4 a-d 1.5 a 1.0 a 1.0 a 3.7 c 4.7 b Ildeunggongshin 15 1.4 a-d 1.6 a 1.0 a 1.0 a 4.0 d 5.0 d Geumnara 14-15 1.5 a-d 2.9 e-i 1.0 a 1.0 a 3.9 d 4.7 bc PR-Taepyeong 15 3.3 h-k 3.7 j-m 1.0 a 1.0 a 4.0 d 5.0 d Yeokganghongjanggun 13 3.3 h-k 3.6 i-m 1.0 a 1.1 ab 4.0 d 5.0 d Ildangbaek 14-15 1.0 a 1.4 a 1.0 a 1.1 ab 4.0 d 5.0 d HNX14460242 15 1.4 a-d 3.2 f-j 1.0 a 1.1 ab 4.0 d 5.0 d Gangryeokdaetong 15 1.9 a-e 3.3 g-k 1.0 a 1.1 a-c 4.0 d 5.0 d Dokyacheongcheong 15 2.0 b-f 2.7 d-h 1.0 a 1.1 ab 4.0 d 4.7 bc Powerspeed 11-12 2.3 d-g 3.4 h-l 1.0 a 1.1 ab 4.0 d 5.0 d Cheonnyeonmannyeon 15 2.9 f-i 3.2 f-j 1.0 a 1.1 ab 4.0 d 5.0 d PR-Sangsaeng 15 3.5 h-k 4.4 n-q 1.0 a 1.1 a-c 4.0 d 5.0 d Josaenggeomul 15 3.6 i-k 4.1 l-o 1.0 a 1.2 a-d 4.0 d 5.0 d Daegeonwang 15 1.3 a-c 1.9 a-c 1.0 a 1.3 a-d 2.3 b 4.7 bc Ilinja 15 1.5 a-d 2.7 d-g 1.0 a 1.3 a-d 4.0 d 5.0 d N6510 15 2.1 c-f 3.4 h-l 1.0 a 1.3 a-d 4.0 d 5.0 d Seonguja 15 2.8 f-i 2.5 b-e 1.0 a 1.3 a-d 4.0 d 5.0 d Giripbaksu 15 2.0 b-f 3.3 g-k 1.0 a 1.4 a-d 4.0 d 4.9 cd Muhanjilju 15 2.2 c-f 2.5 c-f 1.0 a 1.5 a-d 4.0 d 5.0 d PR-Daechon 15 2.7 e-h 3.6 i-m 1.0 a 1.6 b-d 4.0 d 5.0 d PR-Manse 15 3.2 h-k 4.3 m-p 1.1 a 1.3 a-d 4.0 d 5.0 d PR-Bultina 15 2.0 b-f 4.0 k-o 1.1 a 1.3 a-d 4.0 d 5.0 d PR-Oraetta 15 3.2 h-k 3.7 j-n 1.1 a 1.3 a-d 4.0 d 5.0 d Gangryeokjosenggeon 15 3.5 h-k 4.7 o-q 1.1 a 1.7 d 4.0 d 5.0 d PR-Sadaecheonwang 13 3.1 g-j 3.9 j-n 1.3 ab 1.5 a-d 4.0 d 5.0 d PR-Hongdukkae 12-14 3.8 jk 4.3 m-p 1.3 ab 1.7 cd 4.0 d 5.0 d PR-Galmuri 15 4.0 k 4.9 pq 1.3 ab 2.2 e 4.0 d 5.0 d Mansahyeongtong 13-15 4.0 k 5.0 q 1.6 bc 2.8 f 4.0 d 5.0 d Taesan 13-14 2.8 f-i 3.8 j-n 1.8 c 2.8 f 4.0 d 5.0 d Cheongyang 8-15 4.0 k 5.0 q 2.5 d 4.3 g 4.0 d 5.0 d Geumtap 9-15 4.0 k 5.0 q 2.6 de 4.3 g 4.0 d 5.0 d Nokgwang 9-15 3.8 jk 4.9 pq 3.0 e 4.3 g 3.8 cd 4.9 cd x Mean separation within columns by Duncan's multiple range test, P 0.05. y 1 = No root rot observed; 2 = about 25% root rot; 3 = about 50% root rot ; 4 = about 75% root rot; 5 = complete root rot. z 1 = No disease symptoms observed; 2 = necrotic lesion at base of stem, yet still surviving; 3 = wilting; 4 = dead and dried. 32 t 3 t 3 (,, ) ³ w» w (Table 4). w w t, ³, t ³ y z w (Table 5). 3 ³ (Virulence) j ƒ. p Pc005( ) ³ t kk wš û t. w Pc003( )

ü q š t w 145 Table 5. Analysis of variancegfor 2010 experiment for testing resistance of 36 commercial cultivars to 3 isolates of Phytophthora capsici Plant part Source of variation df SS MS Fs Isolate (I) 2 1828.6 914.3 1941.6** Stem Cultivar (C) 35 439.95 12.6 0026.7** I C interaction 70 386.72 5.5 0011.7** Strain (I) 2 2964.2 1482.1 4019.5** Root Cultivar (C) 35 598.2 17.1 0046.4** I C interaction 70 438.1 6.3 0017.0** ³ Pc002( ) ³ w. Pc005( )>Pc002 ( )>Pc003( ) ³ ùkû. t t Pc005( ) ³ û t, Pc003( ) ³ 2009 w w ù t ƒ ƒ w š, Pc002( ) ³ Pc003( ) ù t ƒ ƒ w. Pc002( ) Pc003 ( ) ³ w ww kk, PR-, š, m,, œ, ù, w ùkû. t w ³ w Table 5 t ³ y š ùkû. ³ t w w, ƒ w ƒ. Table 4 PR ks, y, PR-, Pc003( ) ³ ù Pc002( ) ³ ùkü. t ³ p wš ùkü. ƒ t w ³ w w w t w w,» ³ w t w ƒ w v w (Foster Hausbeck, 2010). p w š t ³ w y w w v w. Pc005( ) ³ w ³ xwš ³ t w ƒ w w wš. š ³ š y 2 k (Heterothallic) x(mating type) A1 A2 ƒ ³ ù»»(oogonia)»(antheridia) x w» ù (n) š»» w ùs x w 2 (2n) z w (Erwin Ribeiro, 1996). ü x A1 A2ƒ swš (½, 1988) s k ³ w w y ù w t w yw». š ³(Phytophthora capsici) t p l ƒ w. š ³ w Polach Webster(1972) š, m m, ƒ y m w 23 P. capsici ³» w ƒ š m w (pathogenicity) 14 m(strain) w. ù ü ³ (virulence) jù t ³ p, y (Yang, 1989). ù Hwang (1996) ü ³ w š p šw, Oelke Bosland(2003) w ³ sww 10 ³ CM334 w m t w 9 race w. Glosier (2007) CM334, PI201234 w š m t 34 ³ w 14 x, (race) w. Sy (2008) CM334 Early Jalapeno l w wüy (Recombinant inbred lines, RILs) 17 ³ w 13 w, p w w (Monroy-Barbosa Bosland, 2008). š ³ w š ù ¾ mw y. w y w t ³ w w ƒ ƒ š t w. š w

146 ½ Á«k Áy Á Á Á xá½x PI163192, PI201232, PI201234 AC2258(=Mexican pepper Line 29 ), CM334 (Criollo de Morales 334) (Bosland Lindsey, 1991;, 1985; y ½, 1997; ½ 1986, 1988; Kimble Grogan, 1960), p CM334 ¾ w ³ š w ùkü (Candole, 2010; Foster Hausbeck, 2010; Glosier, 2007; Oelke Bosland, 2003). ü w w w (y ½, 2002). š w w š» w ù(smith, 1967; Gil Ortega, 1990, 1991, 1992; Walker Bosland, 1999) w š (Thabuis, 2004a). (Quantitative trait loci, QTL) ƒ š t w w t w» w y (Bartual, 1991; Palloix, 1990), p QTL mapping j(molecular marker) w z š (Lefevre Palloix, 1996; Thabuis, 2004b). ù w ƒ e w w ³ w ùkü t w v w, pyw w ³ w w w v w. w, s (CMS) w z w QTL š w w» ƒ (y ½, 2002) y w» w (GMS) w w mw w. 2008 l 2010 ü q š š t w sƒw p 2010 t š w 3 ³ x t w w t ³ p w. ü q t w t s. p t w w y PR t w û ù t t k ƒ ƒ, ƒ t t w v w ùkû. 2008-2009 x t 3 ³ w Pc005( ) ³ t kk w t ùkü. Pc003( ) ³ Pc002( ) ³ ƒ w ùkû, t ³ p ƒ t ³ w ùk û ù ³ ùkû. ³ t ³ p w ƒ t w w. py ( y PJ007111201009). š x Bartual, R., Carbonell, E. A., Marsal, J. I., Tello, J. C. and Campos, T. 1991. Gene action in the resistance of peppers (Capsicum annuum) to Phytophthora stem blight (Phytothora capsici L.). Euphytica 54: 195-200. Bosland, P. W. and Lindsey, D. L. 1991. A seedling screen for Phytophthora root rot of pepper, Capsicum annuum. Plant Dis. 75: 1048-1050. Candole, B. L., Conner, P. J. and Ji, P. 2010. Screening Capsicum annuum accessions for resistance to six isolates of Phytophthora capsici. HortSci. 45: 254-259., Ÿ, ¼,,. 1985. w w tš. ( ) 27: 6-11. Erwin, D. C. and Ribeiro, O. K. 1996. Phytophthora diseases worldwide. APS Press. Foster, J. M. and Hausbeck, M. K. 2010. Resistance of pepper to Phytophthora crown, root and fruit rot is affected by isolate virulence. Plant Dis. 94: 24-30. Gil Ortega, R., Palazon Espanol, C. and Cuartero Zueco, J. 1990. Genetics of resistance to Phytophthora capsici in the Mexican pepper Line 29. Bulletin OEPP/EPPO Bulletin 20: 117-112. Gil Ortega, R., Palazon Espanol, C. and Cuartero Zueco, J. 1991. Genetics of resistance to Phytophthora capsici in the pepper line SCM-334. Plant Breeding 107: 50-55. Gil Ortega, R., Palazon Espanol, C. and Cuatero Zueco, J. 1992. Genetic relationships among four pepper genotypes resistant to Phytophthora capsici. Plant Breeding 108: 118-125. Glosier, B. R., Ogundiwin, E. A., Sidhu, G. S., Sischo, D. R. and Prince, J. P. 2007. A differential series of pepper (Capsicum

ü q š t w 147 annuum) lines delineates fourteen physiological races of Phytophthora capsici. Euphytica DOI 10.1007/s10681-007- 9532-1. Hwang, B. K., Kim, Y. J. and Kim, C. H. 1996. Differential interactions of Phytophthora capsici isolates with pepper genotypes at various plant growth stages. Eur. J. Plant Pathol. 102: 311-316. y, ½. 1997. š w m s wü x. w 38: 684-687. y, ½. 2002. š w s z. w 43: 143-150. ½. 1986. š m w. w 27: 11-14. ½. 1988. š w m w m p. w 29: 247-252. ½,. 1992. w š w w» w». w 33: 312-317. ½,,. 1975. š t w w. w 16: 214-219. Kim, B. S., Hwang, H. S., Kim, J. Y. and Han, J. H. 2001. Additional sources of resistance to Phytophthora blight in pepper. Jour. Kor. Soc. Hort. Sci. 42: 233-237. ½,, ½. 1996. w š w w» w z š. w 37: 5-11 ½, ky, ³,. 1988. š ³ (Phytophthora capsici Leonian) x s. w³ 16: 60-63. Kimble, K. A. and Grogen, R. G. 1960. Resistance to Phytophthora root rot pepper. Plant Dis. Rep. 44: 872-873. Lefebvre, V. and Palloix, A. 1996. Both epistatic and additive effects of QTLs are involved in polygenic induced resistance to disease: a case study, the interaction pepper-phytophthora capsici Leonian. Theor. Appl. Genet. 93: 503-511. Monroy-Barbosa, A. and Bosland, P. W. 2008. Genetic Analysis of Phytophthora root rot race-specific resistance in Chile pepper. J. Amer. Soc. Hort. Sci. 133: 825-829. Oelke, L. M. and Bosland, P. W. 2003. Differentiation of racespecific resistance to Phytophthora root rot and foliar blight in Capsicum annuum. J. Amer. Soc. Hort. Sci. 128: 213-218. Palloix, A., Daubeze, A. M., Phaly, T. and Pochard, E. 1990. Breeding Transgressive lines of pepper for resistance to Phytophthora capsici in a recurrent selection system. Euphytica 51: 141-150. Polach, F. J. and Webster, R. K. 1971. Identification of strains and inheritance of pathogenicity in Phytophthora capsici. Phytopathology 62: 20-26. Smith, P. G., Kimble, K. A., Gregan, R. G. and Millett, A. H. 1967. Inheritance of resistance in pepper to Phytophthora root rot. Phytopathology 57: 377-379. Sy, O., Steiner, R. and Bosland, P. W. 2008. Recombinant inbred line differential identifies race-specific resistance to Phytophthora root rot in Capsicum annuum. Phytopathology 98: 867-870. Thabuis, A., Lefebvre, V., Bernard, G., Daubeze, A. M., Phaly, T., Pochard, E. and Palloix, A. 2004a. Phenotypic and molecular evaluation of a recurrent selection program for a polygenic resistance to Phytophthora capsici in pepper, Theor. Appl. Genet. 109: 342-351. Thabuis, A., Palloix, A., Servin, B., Daubeze, A. M., Signoret, P., Hospital, F. and Lefebvre, V. 2004b. Marker-assisted introgression of 4 Phytophthora capsici resistance QTL alleles into a bell pepper line: validation of additive and epistatic effects. Mol. Breed. 14: 9-20. Walker, S. J. and Bosland, P. W. 1999. Inheritance of Phytophthora root rot and foliar blight resistance in pepper. J. Amer. Soc. Hort. Sci. 124: 14-18. Yang, S. S., Sung, N. K., Choi, D. I. and Kim, C. H. 1989. Pathogenic variation of Phytophthora capsici Leonian on redpepper in Korea. Korean J. Plant Pathol. 5: 370-376.