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Euphytica (2006) 149: 97–103 DOI: 10.1007/S10681-005-9057-4

 C

Springer 2006

Evaluation of resistance sources and inheritance of resistance in kidney bean to Indian virulences of Colletotrichum lindemuthianum Evaluation of resistance in bean to anthracnose Anju Pathania1,# , P.N. Sharma1,∗ , O.P. Sharma1 , R.K. Chahota2 , Bilal Ahmad1 & P. Sharma1 1

Molecular Plant Pathology Laboratory, Department of Plant Pathology, CSK HP Agricultural University, Palampur-176 062 (H.P.) India (# Research Associate, Plant Breeding); 2 Assistant Plant Breeder, Department of Plant Breeding & Genetics, CSK HP Agricultural University, Palampur - 176 062 (H.P.) India (∗ author for correspondence: e-mail: [email protected])

Received 31 January 2005; accepted 15 November 2005

Key words: Bean anthracnose, Colletotrichum lindemuthianum, common bean, Phaseolus vulgaris L., resistance

Summary Forty nine common bean lines comprising of exotic accessions and locally grown cultivars evaluated against Colletotrichum lindemuthianum exhibited differential resistance to its races in Himachal Pradesh, a north-western Himalayan state of India. Some exotic accessions like G 2333, Cornell 49242, PI 207262, Mexique 222, TO, Perry Marrow, Kaboon and Widusa were resistant to more than five Indian races, whereas two Indian accessions KRC-5 and Hans showed resistance to six and four races, respectively. However, nine accessions KRC-8, KR-40, KR-43, KR-81, KR-62-2, KR-90, KR-142, KR-148, and KR-216 were resistant to three races. Race specific resistance has been observed in different bean cultivars. Studies on inheritance of resistance in exotic accession G 2333 and Indian accession, KRC-5 showed that two independent dominant genes conferred resistance in G 2333 to race 3 and 515 and a single dominant gene controlled resistance in KRC-5 to race 775, indicating resistance from these sources is easily transferable to the locally adapted susceptible cultivars.

Introduction Common bean (Phaseolus vulgaris L.), occupies premier place among grain legumes in the world including India where it is locally called as ‘Rajmash’ (Gepts & Debouck, 1991; Tu, 1992; Sharma et al., 1994). It is a native of central and south America and is morphologically highly variable. Common bean is adapted to a wide range of environments under diverse agroeco systems (Popelka et al., 2004). In India, beans are grown in an area of about nine million hectares with an annual production of three million tonnes (FAO, 2000). Similarly, common beans grown all over Himachal Pradesh, a north- western Himalayan state of India also possess extreme morphological variability, though not fully exploited as compared to Andean and

Mesoamerican bean gene pools (Sharma et al., 1994; Chaudhury, 1997). Such a diversity in Indian beans cultivated in north- western Himalayan region could be due to its location which is close to the secondary centre of origin, western China (Tibet) (Vavilov, 1951; Singh, 2001). Anthracnose caused by Colletotrichum lindemuthianum (Sacc. & Magn.) Bri. & Cav., is the major constraint for quality bean production in Himachal Pradesh. The disease is wide spread in sub temperate to temperate region where cool and humid environment favours the disease development resulting in significant yield losses (Chakrabarty et al., 1985; Shao and Teri, 1985; Pastor-Corrales and Tu, 1989; Fernandez et al., 2000; Sharma et al., 2004). Like elsewhere the pathogen has been found to possess high variability in

98 Himachal Pradesh. Variation in the pathogen populations has been studied on bean differentials and through DNA fingerprinting (Sharma et al., 1999; Sharma et al., 2003). Due to highly variable nature of the pathogen, there has been frequent break down of resistance in different parts of the world (Fouilloux, 1976; Menezes and Dianese, 1988; Kelly et al., 1994; Pastor- Corrales et al., 1994, 1995; CIAT, 1995; Mahuku et al., 2002 and Mahuku & Riascos, 2004). Majority of the recommended and locally adapted bean cultivars in Himachal Pradesh are susceptible to different races of pathogen (Kumar et al., 1997 and Sharma et al., 1999). The success of a particular cultivar depends upon its agronomic and quality traits, however, most of the bean accessions resistant to different races of pathogen have long duration and indeterminate growth, a trait impalatable to the farmers. Resistance to disease has been studied systematically in USA and Europe resulting in identification of resistance sources among Andean and Mesoamerican gene pools of common beans and molecular markers linked to six dominant genes have been identified (Schwartz et al., 1982; Balardin et al., 1990; Young and Kelly, 1996, 1997; Young et al., 1998; Geffroy et al., 1999; Mendoza et al., 2000; Vallejo and Kelly, 2001, 2002). Resistance has been found to be conditioned by nine major genes Co-1 to Co- 8 and Co- 10 and one recessive gene Co-8 (Kelly and Vallejo, 2004). No systematic study has been conducted on resistance in common beans and its pathogen in the Indian subcontinent except few reports emanating from this laboratory depicting variation in bean anthracnose pathogen and presence of resistance in some locally grown cultivars (Sharma et al., 1993; Sharma et al., 1999; Sharma et al., 2004). During resistance breeding, it is necessary to understand the nature of resistance, its suitability in resistance breeding and its temporal and spatial deployment. This paper reports the results of evaluation common bean accessions and nature of resistance in an exotic accession G-2333 and in an indigenous one KRC-5 to Indian virulences of C. lindemuthianum.

Materials and methods Race cultures, preparation of inoculum and method of inoculation Monosporic cultures of ten races of C. lindemuthianum maintained in this laboratory were used in the studies. The virulence of each race was confirmed

by inoculating a set of 12 common bean differentials (Pastor- Corrales, 1991). These races belong to the Mesoamerican race group based on the description of Mahuku and Riascos (2004). Spore suspension was made in sterilized water from seven days old sporulating culture of each race. The suspension was filtered through double layered muslin cloth and inoculum load was adjusted to 2.1 × 106 spores ml−1 with the help of a heamocytometer. Germinated seed dip method (Champion et al., 1973) was used for inoculation of the test plants. Three days old germinating five seeds of each test variety were dip-inoculated in spore suspension for 3–5 minutes after removing the seed coats. The inoculated seeds were sown in sterilized sand in iron trays and were kept at 22 ± 1◦ C with 90% relative humidity and 12 h photoperiod in a growth chamber (Saveer India Ltd.) up to 12 days. The disease reaction was scored after six and twelve days of inoculation following 0–5 scale (Drijfhout & Davis, 1986), where 0 = no disease; 1 = pin point lesions; 2 = small lesions, not sunken; 3 = large sunken lesions; 4 = large, deep lesions up to stem centre and 5 = seedlings killed by the pathogen. Plants scoring reaction type of 0, 1, 2 were graded as resistant while those scoring 3, 4, and 5 were graded as susceptible. The reaction of all the differentials to each isolate was evaluated twice. Evaluation of germplasm Forty nine common bean genotypes comprising of land races, recommended varieties and exotic accessions, were evaluated for resistance to ten races of C. lindemuthianum under artificial epiphytotic conditions (Table 1).The seeds of land races and recommended varieties were obtained from Mountain Agriculture and Extension Centre, Sangla, CSK HPAU, Palampur, whereas exotic accessions were procured from CIAT, Cali, Colombia. Land races comprised of single plant selections from the germplasm collected from dry temperate region of Himachal Pradesh. Five seeds of each line were used for inoculation purpose. The resistance was confirmed twice using five seeds per test. Procedure for resistance evaluation was adopted as described in the previous section. Inheritance of resistance Indian land race KRC-5 and an exotic accession G2333 possessing resistance to different races of C. lindemuthianum were used as resistant and locally adapted cultivars Jawala and Kanchan as susceptible

99 Table 1. Reaction of kidney bean germplasm to different races of Colletotrichum lindemuthianum Reaction Races Resistant

Susceptible

3

KR-142, KR-62-2, KR-90, Hans, KRC-5, G 2333, Perry Marrow, Cornell 49242, Widusa, Kaboon, Mexique 222, PI 207262, TO, TU

115

KRC-8, G 2333, Perry Marrow, Cornell 49242, Kaboon, PI 207262, TO, TU

513

KR-220, KR-57, KR-256, KR-43, KR-141-A, SR-6, KR-45, KR-216, KR-178, KR-160, KR-243, KR-256-3, KR-172, KR-293, KR-257, KR-183, KR-175-3, KR-40, KR-81, KR-139, KR-46, SR-1, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-94, KR-52-2, KR-148, EC-57080, Jawala, Kanchan, KRC-8, Michigan dark red kidney KR-220, KR-57, KR-256, KR-43, KR-141-A, SR-6, KR-45, KR-142, KR-216, KR-178, KR-160, KR-243, KR-62-2, KR-256-3, KR-172, KR-293, KR-257, KR-183, KR-175-3, KR-40, KR-81, KR-139, KR-46, SR-1, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-90, KR-94, KR-52-2, KR-148, Hans, EC-57080, Jawala, Kanchan, KRC-5, Michigan dark red kidney, Widusa, Mexique 222 KR-220, KR-57, KR-256, KR-141-A, KR-45, KR-142, KR-216, KR-178, KR-160, KR-62-2, KR-256-3, KR-172, KR-293, KR-257, KR-183, KR-175-3, KR-46, SR-1, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-90, KR-94, KR-52-2, Hans, EC-57080, TU

KR-43, SR-6, KR-243, KR-40, KR-81, KR-139, KR-148, Jawala, Kanchan, KRC-5, KRC-8, G 2333, Michigan dark red kidney, Perry Marrow, Cornell 49242, Widusa, Kaboon, Mexique 222, PI 207262, TO KR-45, KR-142, KR-216, KR-62-2, KR-220, KR-57, KR-256, KR-43, KR-141-A, SR-6, KR-178, KR-160, KR-243, KR-183, KR-303, KR-90, Hans, KR-256-3, KR-172, KR-293, KR-257, KR-175-3, KR-40, KR-81, KR-139, EC-57080, G 2333, Perry Marrow, KR-46, SR-1, KR-155-3, KR-65-2, KR-9, KR-39, KR-94, KR-52-2, KR-148, Cornell 49242, Widusa, Kaboon, Mexique Jawala, Kanchan, KRC-5, KRC-8, Michigan dark red kidney, TU 222, PI 207262, TO KR-43, SR-6, KR-40, KR-81, KR-139, KR-220, KR-57, KR-256, KR-141-A, KR-45, KR-142, KR-216, KR-178, SR-1, Jawala, Kanchan, KRC-5, KRC-8, KR-160, KR-243, KR-62-2, KR-256-3, KR-172, KR-293, KR-257, KR-183, G 2333, Michigan dark red kidney, Perry KR-175-3, KR-46, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-90, Marrow, Cornell 49242, Kaboon, KR-94, KR-52-2, KR-148, Hans, EC-57080, Widusa, TU Mexique 222, PI 207262, TO KR-43, KR-243, KR-40, KR-81, KRC-5, G KR-220, KR-57, KR-256, KR-141-A, SR-6, KR-45, KR-142, KR-216, KR-178, 2333, Kaboon, Mexique 222, PI 207262, KR-160, KR-62-2, KR-256-3, KR-172, KR-293, KR-257, KR-183, KR-175-3, TO KR-139, KR-46, SR-1, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-90, KR-94, KR-52-2, KR-148, Hans, EC-57080, Jawala, Kanchan, KRC-8, Michigan dark red kidney, Perry Marrow, Cornell 49242, Widusa, TU

515

529

537

615

631

775

935

KR-178, KR-148, G 2333, Cornell 49242, Widusa, PI 207262, TO

KR-220, KR-57, KR-256, KR-43, KR-141-A, SR-6, KR-45, KR-142, KR-216, KR-160, KR-243, KR-62-2, KR-256-3, KR-172, KR-293, KR-257, KR-183, KR-175-3, KR-40, KR-81, KR-139, KR-46, SR-1, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-90, KR-94, KR-52-2, Hans, EC-57080, Jawala, Kanchan, KRC-5, KRC-8, Michigan dark red kidney, Perry Marrow, Kaboon, Mexique 222, TU KR-148, G 2333, Cornell 49242, PI 207262, KR-220, KR-57, KR-256, KR-43, KR-141-A, SR-6, KR-45, KR-142, KR-216, TO KR-178, KR-160, KR-243, KR-62-2, KR-256-3, KR-172, KR-293, KR-257, KR-183, KR-175-3, KR-40, KR-81, KR-139, KR-46, SR-1, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-90, KR-94, KR-52-2, Hans, EC-57080, Jawala, Kanchan, KRC-5, KRC-8, Michigan dark red kidney, Perry Marrow, Widusa, Kaboon, Mexique 222, TU KR-45, KR-142, KR-216, KR-178, KR-220, KR-57, KR-256, KR-43, KR-141-A, SR-6, KR-160, KR-243, KR-256-3, KR-62-2, KR-90, Hans, EC-57080, KR-172, KR-293, KR-257, KR-183, KR-175-3, KR-40, KR-81, KR-139, KRC-5, G 2333, Cornell 49242, Widusa, KR-46, SR-1, KR-155-3, KR-303, KR-65-2, KR-9, KR-39, KR-94, KR-52-2, Kaboon, Mexique 222, PI 207262, KR-148, Jawala, Kanchan, KRC-8, Michigan dark red kidney, Perry Marrow, TO, TU KR-216, KR-172, KR-183, KR-303, Hans, KR-220, KR-57, KR-256, KR-43, KR-141-A, SR-6, KR-45, KR-142, KR-178, EC-57080, KRC-5, G 2333, Cornell KR-160, KR-243, KR-62-2, KR-256-3, KR-293, KR-257, KR-175-3, KR-40, 49242, Widusa, Mexique 222 KR-81, KR-139, KR-46, SR-1, KR-155-3, KR-65-2, KR-9, KR-39, KR-90, KR-94, KR-52-2, KR-148, Jawala, Kanchan, KRC-8, Michigan dark red kidney, Perry Marrow, Kaboon, PI 207262, TO, TU

100 parents. G-2333, a Mesoamerican accession has small and maroon coloured seed, indeterminate growth habit whereas KRC-5 is large seeded violet coloured indeterminate and semi climbing type. All the resistant accessions are late in flowering and maturity by 15– 20 days as compared to susceptible cultivars. Sowing of both resistant and susceptible parents was done at fortnight intervals from last week of April, 2002 to June, 2002, to prolong the flowering period so that maximum crosses could be attempted. Sowing of resistant parents was done 15–20 days earlier in order to coincide the flowering period of both parents. Resistant parents were crossed with susceptible ones using manual emasculation and pollination. In each cross five to six hundred flower buds were used to obtain sufficient F1 seed as the success rate of hybridization in beans is quite low. All hybridization and seed production work was carried out under isolated conditions at university Mountain Agriculture Research & Extension Centre, Sangla (Kinnaur) situated at 2600 meters above mean sea level. This place represents the main bean growing areas and is ideally suited to breeding work. The cool and humid climate of the area results in anthracnose epidemics. The reaction pattern of parents was evaluated in the growth chamber at seedling stage using seedling dip method (Champion et al., 1973) and under field conditions at adult plant stage using spray inoculation method, where as F1 and F2 generations were evaluated only at seedling stage as the parents did not exhibit differential reaction at adult plant stage. Since reactions were discrete, data were classified into class frequencies and tested for goodness of fit with chi-square test.

KRC-8, KR-40, KR-43, KR-81, KR-62-2, KR-90, KR142, KR-148, and KR-216 to three races. However, KRC-8 was found highly resistant under field conditions. Sixteen indigenous accessions were susceptible to all the races of the pathogen. Remaining cultivars possessed race specific resistance. Perusal of the literature on bean resistance to C. lindemuthianum in Europe and USA revealed numerous sources of resistance (Balardin et al., 1990; Schwartz et al., 1982), however, few such data with respect to India (Sharma et al., 1999) are available. G 2333 is resistant to almost all the European and American isolates (Kelly et al., 1994; Pastor-Corrales et al., 1994). However, the breakdown of resistance in G 2333 has been reported against races from Costa Rica, Mexico and Argentina (CIAT, 1995; Mahuku et al., 2002). Cornell 49242 having Are gene was found susceptible to race 537 confirming the report of its breakdown by Fouilloux (1976) and Kelly et al. (1994). Similarly breakdown of resistance in Mexique 222, TO and TU has also been reported against Middle Latin and American races of C. lindemuthianum (PastorCorrales et al., 1994, 1995). Resistance in different locally grown and exotic beans has also been reported by Kumar et al. (1997) and Sharma et al. (1999). Since pathotypes used in this study belong to Mesoamerican race group which infects cultivars of both Andean and Mesoamerican gene pool, sources of resistance found in this study could be related to either of the gene pool since information about the common origin of Indian common beans and Andean and Mesoamerican gene pools is not available. Inheritance of resistance

Results and discussion Evaluation of resistance Evaluation of different common bean genotypes against ten races of C. lindemuthianum (Table 1) revealed that lines/accessions widely used in Europe, America and Brazil exhibited resistance to majority of Indian races. Accession G 2333 was found highly resistant to all the races followed by Cornell 49242 and PI 207262 (Table 1), remaining exotic accessions exhibited resistance to five or more races. Exotic accessions Michigan dark red kidney and TU were found susceptible to almost all races. Among indigenous germplasm two accessions KRC-5 and Hans were found resistant to six and four races, respectively and nine accessions

Reactions of different test cultivars to various races of C. lindemuthianum are presented in Table 2.The F1 plants of different crosses exhibited resistant reaction indicating resistance to be dominant. The observed and expected frequencies of resistant and susceptible reaction of parents, F1 and F2 populations are also presented in the Table 2. The chi square (χ 2 ) values showed a good fit for segregation ratio of 15 resistant to 1 susceptible in F2 populations of Jawala × G 2333 and Kanchan x G 2333 and 3 resistant to 1 susceptible in Jawala x KRC-5 and Kanchan × KRC-5. Segregation ratio of 15 resistant to 1 susceptible in F2 populations of Jawala × G 2333 and Kanchan × G 2333 indicates that the resistance in G 2333 to Indian race 515 is controlled by two independent dominant genes with equal effects. G 2333 has been reported

101 Table 2. Segregation for resistance in KRC-5 and G 2333 to races 3, 515 and 775 of Colletotrichum lindemuthianum Number of seedlings Parents/Crosses Race 775 Jawala KRC-5 Jawala × KRC-5 Kanchan KRC-5 Kanchan × KRC-5 Race 3 Jawala G 2333 Jawala × G 2333 Kanchan G 2333 Kanchan × G 2333 Race 515 Jawala G 2333 Jawala × G 2333 Kanchan G 2333 Kanchan × G 2333

Generation

Resistant

Susceptible

Expected ratio

χ2

P ≤ 0.05

P1 P2 F1 F2 P1 P2 F1 F2

– 25 10 149 – 25 12 164

25 – – 51 25 – – 56

– – – 3:1 – – – 3:1

– – – 0.03 – – – 0.02

– – – 0.7–0.9 – – – 0.7–0.9

P1 P2 F1 F2 P1 P2 F1 F2

– 15 10 119 – 15 10 115

25 – – 7 25 – – 8

– – – 15:1 – – – 15:1

– – – 0.10 – – – 0.01

– – – 0.7–0.9 – – – 0.9–0.95

P1 P2 F1 F2 P1 P2 F1 F2

– 15 10 160 – 15 10 170

25 – – 11 25 – – 10

– – – 15:1 – – – 15:1

– – – 0.01 – – – 0.15

– – – 0.9–0.95 – – – 0.7

P 1: Parent 1; P 2: Parent 2.

to possess three genes Co-42 , Co-5 and Co-7 (CIAT, 1995; Kelly and Vallejo, 2004). Co-5 gene is present in differential cultivar TU (Kelly and Vallejo, 2004) on which race 515 has been found to be virulent indicating the susceptibility of Co5 gene also present in cultivar G 2333. Thus, segregation ratio of 15:1 seems to be due to the involvement of two genes. Present studies confirms the report of Pastor-Corrales et al. (1994) who found accession G 2333 to possess duplicate dominant genes conferring resistance to Colombian isolate (race 521). Young and Kelly (1996) explained that the duplicate type of gene action in G 2333 as reported by Pastor- Corrales et al. (1994) has been due to virulence of race 521 to differential cultivar TU. Similarly, Cardenas et al. (1964), Muhalet et al. (1981) and Peloso et al. (1989a) have also reported that duplicate genes were responsible for anthracnose resistance in some bean crosses. Similar segregation ratio of 15 resistant to 1 susceptible in F2 population of Jawala × G 2333 and Kanchan

× G 2333 was noticed against race 3 confirming the involvement of two independent dominant genes in G 2333. However, the presence of three genes (Co-42 , Co-5 and Co-7) in G 2333 possessing resistance to different races could not be established in this crossrace combination as race 3 is avirulent on all the three genes present in TO, TU, and G 2333. Mahuku and Riascos (2004) report the susceptibility of G 2333 to races 3481 and 3545 from Costa Rica and Argentina which indicate that these races are virulent to Co-42 , Co-5 and Co-7 genes, however, TU with Co-5 gene is resistant to these races indicating interaction between Co-5 gene, Co-42 and Co-7 genes. The segregation ratio of 3 resistant to 1 susceptible in F2 populations of Jawala × KRC-5 and Kanchan × KRC-5 reveals the presence of single dominant gene for resistance in KRC-5 against race 775. Monogenic dominant resistance for anthracnose in common bean has also been reported by many workers (Mastenbrock, 1960; Cardenas et al., 1964; Bannerot et al., 1971;

102 Muhalet et al., 1981; Peloso et al., 1989; TU, 1992). Monogenic dominant nature of anthracnose resistance in KRC-5 is also confirmed by our earlier findings (Sharma et al., 2000). However, Rudorf (1961) reported monogenic recessive resistance to race Beta of the pathogen and digenic recessive to alpha, gamma and delta races. The present studies establish that the resistance in KRC-5 and G 2333 to different races of the pathogen is governed by dominant genes. The monogenic resistance observed in KRC-5 though, conferred resistance to six races of the pathogen, although variability in C. lindemuthianum points to its vulnerability. The resistance in common bean accessions can be exploited in disease resistance breeding programme through gene pyramiding. However, more Indian germplasm need to be exploited for new resistance genes as wide diversity exists in common beans of north-western Himalayan region (Chaudhary, 1997). Dominant nature of resistance in accession G 2333 and local land race KRC-5 also revealed that resistance is transferable to locally adapted cultivars following any detection method provided both dominant alleles are transferred.

Acknowledgments The authors are thankful to Indian Council of Agricultural Research for financial support in the form of Adhoc research project; Associate Director, Mountain agriculture Research and extension Centre, Sangla (Kinnaur) H.P. for providing facilities for crossing work and generation advancement; reviewers and Prof. R.P. Kaushal, for critical review of this manuscript.

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