Introduction
Rice sheath blight caused by soil borne necrotrophic fungus Rhizoctonia solani is one of the major rice (Oryza sativa L.) disease (Lee and Rush, Reference Lee and Rush1983; Rush and Lee, Reference Rush, Lee, Webster and Gunnell1992). The yield losses due to sheath blight are recorded up to 69% under the intensive crop management and favourable environmental conditions (Sivalingam et al., Reference Sivalingam, Vishwakarma and Singh2006). The pathogen has a wide host range and necrotrophic nature. Moreover, evaluation of sheath blight resistance in rice fields is very difficult because resistance is largely affected by morphological characteristics such as plant height (PH), plant type, tillering, heading date and ecological aspects such as water and N fertilization (Hashiba et al., Reference Hashiba, Uchiyamada and Kimura1981; Groth and Nowick Reference Groth and Nowick1992). As a consequence, no source of complete genetic resistance or immunity to sheath blight is known in O. sativa (Chen et al., Reference Chen, Zou, Xu, Tong, Tang, Wang, Jiang, Ning, Tang and Pan2000; Eizenga et al., Reference Eizenga, Lee and Rutger2002; Liu et al., Reference Liu, Chen, Ji and Liu2006; Zhang et al., Reference Zhang, Chen, Lu, Yang and Yu2006). Thus, exploring different germplasm became an international effort for getting resistance to sheath blight (Xie et al., Reference Xie, Linscombe, Rush and Jodari-Karimi1992; Jia et al., Reference Jia, Correa-Victoria, McClung, Zhu, Liu, Wamishe, Xie, Marchetti, Pinson, Rutger and Correll2007; Zuo et al., Reference Zuo, Wang, Chen, Gu, Zhang, Chen, Pan and Pan2009). Wild species germplasm of rice constitute a valuable resource for the improvement of modern rice cultivars in terms of providing resistance/tolerance against biotic and abiotic stresses (Khush et al., Reference Khush, Balacangco and Ogawa1990; Rangel et al., Reference Rangel, Brondani, Rangel and Brondani2008; Zhang and Xie, Reference Zhang, Xie and Wengui2014; Arbelaez et al., Reference Arbelaez, Moreno and Singh2015; Eizenga et al., Reference Eizenga, Neves, Bryant, Agrama and Mackill2016; Ma et al., Reference Ma, Fu, Zhao, Jiang, Zhu, Gu, Xu, Su, Sun and Tan2016; Haritha et al., Reference Haritha, Swamy and Naik2018; Kishor et al., Reference Kishor, Sarao, Bhatia, Neelam, Kaur, Mangat, Brar and Singh2018; Quan et al., Reference Quan, Wang and Hui2018). An investigation was carried out to identify potential sources for resistance against sheath blight in Oryza nivara accessions at Punjab Agricultural University (PAU), Ludhiana during 2015, 2016 and 2017.
Experimental
The plant materials consisted of 67 accessions of O. nivara were originally procured from International Rice Research Institute (IRRI), Manila, Philippines and National Rice Research Institute (NRRI), Cuttack, India. Isolation of Rhizoctonia solani was performed from the susceptible cultivar PR121 showing the sheath blight symptoms. The standard protocol for isolation was used. The maize meal-sand (1:3) medium method was used for inoculation (Lore et al., Reference Lore, Hunjan and Thind2012). Disease assessment was made 21 days after inoculation under field conditions. Three plants from each accession in two replications were screened every year. Relative lesion height (RLH) was calculated using the formula given by Sharma et al. (Reference Sharma, Teng and Olivares1990). Data were analysed using the general linear model procedure in the SAS system Cary, NC). The cluster is reported as unweighted pair group method using arithmetic means based on the Mahalanobis distance algorithm (PAST 3.0).
Results
A total of 67 accessions with susceptible check PR114 were screened during 2015. Out of these, 16 (23.88%) were moderately resistant (MR), 21 (31.34%) were moderately susceptible (MS) and 30 (44.77%) were susceptible (S) (online Supplementary Figure S1). The PH ranged from 44 to 171 cm, lesion height (LH) ranged from 21 to 99 cm, RLH ranged from 18.84 to 97.67% and DS (DS) ranged from 1 to 9 (online Supplementary Table S1). All the accessions were clustered into four groups (M1, M2, M3 and M4) by the Mahalanobis distance on the basis of RLH and DS. The group M2 with 16 accessions showed the MR reaction with the minimum values of LH, RLH and DS, i.e. 34.56 cm, 22.67% and 3 respectively. The group M4 had the maximum values of LH, RLH and DS, i.e. 70.48 cm, 74.84% and 8.2 respectively (online Supplementary Table S2). All the genotypes and genotype × environment interaction had a significant (P < 0.001) effect on all the disease variables. The environment interaction had a significant (P < 0.001) effect on all the disease variables except LH (P > 0.30) (online Supplementary Table S3).
Based on screening during the year 2015, 16 MR accessions were selected and further evaluated during the years 2016 and 2017 (Table 1). The PH ranged from 81 to 158 cm, the LH ranged from 12 to 72 cm, the RLH ranged from 8.33 to 67.28% and the DS ranged from 1 to 9 for both the years. The different disease variables measured were significantly (P < 0.05) correlated. RLH had a positive correlation with LH (0.86) and DS (0.93). The LH showed positive correlation with DS, with a correlation coefficient of 0.81. There was a negative correlation between PH and DS, with a correlation coefficient of −0.24 (online Supplementary Table S4). Sixteen accessions O. nivara were selected and clustered into three groups (M1, M2 and M3) on the basis of different disease variables (Fig. 1). The first group M1 with susceptible check PR114 showed the highest values of LH (63.33 cm), RLH (61.50%) and DS (7.0) (online Supplementary Figure S2).
Discussion
Several attempts were made to identify sources for sheath blight resistance in rice genotypes but only partial resistance was identified to date (Amante et al., Reference Amante, de la Pena, Sitch, Leung and Mew1990; Pan et al., Reference Pan, Rush, Sha, Linscombe, Stetina and Oard1999; Prasad and Eizenga, Reference Prasad and Eizenga2008; Ram et al., Reference Ram, Majumdar, Laha, Ansari, Kar and Mishra2008). We also observed partial resistance in the studied O. nivara accessions. The most probable explanation is the pathogen behaviour like necrotrophic nature, wide host range, soil borne and anastomosis. These characters make the pathogen to invade positively and lesion progression is higher than the resistant level. The other attributes includes significant effects of GXE interaction on disease variables which ultimately resulted in moderate resistance behaviour of studied germplasm. Zeng et al. (Reference Zeng, Shi, Ji, Wen, Liang and Yang2017) studied the environmental effect on disease progression among 169 genotypes and observed a highly significant (P < 0.0001) effect. A positive correlation between RLH and DS was observed. This is due to the fact that the LH is determining factor for the progression of disease and the RLH is directly proportional to LH. Similarly, a negative correlation was explicable between PH and RLH as the RLH is the ratio of LH and PH. Similar relationship among disease variables is also reported by other scientist (Hossain et al., Reference Hossain, Tze, Nadarajah, Jena, Bhuiyan and Ratnam2014; Wen et al., Reference Wen, Zeng, Ji and Yang2015). We identified O. nivara accessions with possible sheath blight resistance and were crossed with the elite cultivars to generate backcrossed derivatives for transferring the sheath blight resistance.
Supplementary material
The supplementary material for this article can be found at https://doi.org/10.1017/S1479262119000315.
Acknowledgements
The Authors are grateful to the ICAR for funding under the Project ‘Incentivizing research in agriculture project-IV-Molecular genetic analysis of resistance/tolerance to different stress (Rice)’ CS/F.No. 16-8/17-IA IV. We are also grateful to IRRI, Philippines and NRRI, Cuttack India for sharing wild species germplasm.
Conflict of interest
The authors declare that they have no conflicts of interest.