Plant breeding

Inheritance of vegetative-stage drought tolerance in interspecific and intraspecific crosses of rice
 
Jimmy Lamo, National Crops Resources Research Institute, P.O. Box 7084, Kampala, Uganda; Pangirayi Tongoona, African Centre for Crop Improvement, University of KwaZulu-Natal, Private Bag X01, Scottsville 3209, KwaZulu-Natal, South Africa; Patrick Okori, Department of Crop Science, Makerere University, P.O. Box 7062, Kampala, Uganda; and John Derera and Mark Laing, African Centre for Crop Improvement, University of KwaZulu-Natal, Private Bag X01, Scottsville 3209, KwaZulu-Natal, South Africa E-mail: lamojim@gmail.com; 204524834@ukzn.ac.za 

      Drought is a major constraint to global upland and rainfed lowland rice production. In sub-Saharan Africa (SSA), there is increasing expansion of rice production from traditional irrigated production areas to rainfed environments where the drought problem is an inherent challenge (Balasubramanian et al 2007). New generations of rice, interspecific rice genotypes, offer potential for use in breeding for improved drought tolerance in the region (Majerus et al 2007), based on their high osmotic potential (Majerus et al 2007), high tillering and leaf-rolling capacity, and high plant vigor (Jones et al 1997). In Uganda, a few new interspecific lines have been released as varieties NERICA 1, NERICA 10, and NERICA 4 (Lamo 2010) for cultivation by farmers. These varieties have varying tolerance for drought stress (Kijima et al 2006). Understanding the genetics of drought tolerance or its component traits is necessary to improve the presently cultivated varieties using interspecific lines. Breeding for drought tolerance is a cost-effective option of managing this constraint (Fukai and Cooper 1995). Secondary traits with high heritability for drought tolerance, including leaf drying, leaf rolling, filled grains, root traits, and water-use efficiency in crosses between Oryza sativa and O. glaberrima, have been been reported (Singh and Mackill 1990, Garrity and O’Toole 1994, Lafitte et al 2003, Fujii et al 2005). Singh and Mackill (1990) found that a single gene was responsible for leaf rolling in rice in O. sativa. This study aimed to determine the heritability estimates for leaf rolling under drought during the vegetative growth stage in crosses between O. sativa and interspecific lines developed through crossing O. sativa and O. glaberrima. In this experiment, seven parents were crossed in pairs detailed in Table 1. In each cross, one parent was intraspecific and the other interspecific. In the experiment, the seven parents, seven F1, and seven F2 populations were planted
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Plant breeding

in a 3 × 7 alpha lattice design with two replications. The parental generations and F1s were planted in five-row and three-column plots, whereas the F2 progenies were planted in five-row and six-column plots. The plants were thinned to one plant per hill. Supplementary irrigation was applied at 20 mm per week. Standard cultural practices were followed. Data on the effect of drought stress at the vegetative growth stage were collected. In this method, irrigation was terminated when about 50% of the plants had attained maximum tillering. The leaf-roll score was taken using a 0–9 scale: 0, healthy; 1, shallow Vshaped leaves; 3, deep V-shaped leaves; 5, fully capped leaves; 7, leaf margin tightly held in U-shape; 9, tightly rolled leaves (Gregorio and Cabuslay 2005). It is a modification of the standard evaluation system for rice developed by IRRI (IRRI 2002).
Table 1. Rice genotypes used in the three experiments.
Genotype no. 002 Set D 012 072 121 147 197 129 Breeding line CT16346-CA-20-M CT16344-CA-9-M CT16317-CA-4-M CK 73 IRAT104 IRAT257 CK73 Type Interspecific Interspecific Interspecific O. sativa O. sativa O. sativa O. sativa Rice parent Female Female Female Female Male Male Male

Heritability was determined for each set of the crosses using regression analyses using offspring regression Y; this involves regressing F2 on F1 progenies and F1 progenies on mid-parents (Falconer and Mackay 1996). The regression model was Y = bx + c, where Y is the relationship between F1 and mid-parents and that between F2 and F1, x is the intercept, and c is a constant. The results are summarized in Table 2. Consistently, heritability estimates were higher for regression of F1 on mid-parents than when F2 was regressed on F1 progenies, except for the cross between parent CT16344-CA-9-M and parent WITA 1. Consistent heritability estimates for both methods were observed for crosses CT16344-CA-9-M/CK 73 and CT16344-CA-9-M/WITA 1. The heritability estimates for regression of F2 on F1 progenies were approximately half of the estimates for regression of F1 on mid-parents for crosses CT16346-CA-20M/IRAT 104 and CT 16346-CA-20-M/121. This finding is consistent with what Efisue et al (2009) detected: high and significant heritability estimates of leaf scores in interspecific rice populations under drought stress. Sirault et al (2007) also found that additive effects were the most important components in the control of drought stress in wheat. This study also noted high heritability for leaf
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roll among populations under drought stress, implying that additive effects are important.    
Table 2. Summary of parental, F1, and F2 means and heritability estimates using F1 on mid-parents and F2 on F1 progenies for leaf-roll scores under drought stress.
Parent 1 Var 1 2 3 4 5 6 7
a

Parent 2 Var 121 129 121 129 147 197 147
b

F1 Mean 5.4 ± 0.9 5.3 ± 1.0 4.6 ± 1.2 4.1 ± 1.1 5.2 ± 1.2 5.1 ± 0.8 5.2 ± 1.1

F2 Mean 4.5 ± 0.8 4.7 ± 0.6 4.6 ± 0.5 4.4 ± 0.5 4.6 ± 0.5 4.6 ± 0.5 4.8 ± 0.5

a

Mean 5.9 ± 1.5 5.9 ± 1.5 4.8 ± 1.3 5.9 ± 1.1 5.6 ± 1.1 4.8 ± 1.3 4.8 ± 1.3

Mean 3.2 ± 1.9 4.0 ± 1.4 3.2 ± 1.9 4.0 ± 1.4 3.9 ± 1.5 3.6 ± 1.4 3.9 ± 1.5

F1 on mid-parents h2 0.57 0.65 0.86 0.61 0.94 0.66 0.86

F2 on F1 h2 0.68 0.65 0.38 0.37 0.47 0.49 0.42

12 12 2 2 72 2 2

Variety used in the cross as female parent: 12 = CT 16344-CA-9-M; 2 = CT 16346-CA-20-M; 72 = CT b 16317-CA-4-M. Variety used in the cross as male parent: 121 = WITA 1; 129 = CT 16346-CA-20-M; 147 = IRAT 104; 197 = IRAT 257. 

 

References
Balasubramanian V, Sié M, Hijmans R, Otsuka K. 2007. Increasing rice production in Africa: challenges and opportunities. Adv. Agron. 94:55-133. Efisue AA, Tongoona P, Derera J, Ubi BE, Oselebe HO. 2009. Genetics of morpho-physiological traits in segregating populations of interspecific hybrid rice under stress and non-stress conditions. J. Crop Improve. 23(4):383-401. Falconer DS, Mackay TFC. 1996. Introduction to quantitative genetics. Essex (England): Longman Scientific and Technical Co. 52 p. Fukai S, Cooper M. 1995. Development of drought-resistant cultivars using physio-morphological traits in rice. Field Crops Res. 40:67-87. Fujii M, Andoh C, Ishihara S. 2005. Drought resistance of NERICA (New Rice for Africa) compared with Oryza sativa L. and millet evaluated by stomatal conductance and soil content. Proceedings of the 4th International Crop Science Conference. Melbourne (Victoria): Australian Society of Agronomy. Garrity DP, O’Toole JC. 1994. Screening rice for drought resistance at the reproductive phase. Field Crops Res. 39:99-110. Gregorio GB, Cabuslay GS. 2005. Breeding for abiotic stress. In: Ashraf M, Harris PJC, eds. Abiotic stresses: plant resistance through breeding and molecular approaches. Coventry (UK): Coventry University. p 525-527. IRRI (International Rice Research Institute). 2002. Field evaluation handbook of the International Rice Research Institute. Manila (Philippines): IRRI. Jones MP, Audebert A, Mande S, Aluko K. 1997. Characterization and utilization of Oryza glaberrima Steud. In: Upland rice breeding. In: Jones MP, Dingkuhn M, Johnson DE, Fagade SO, eds. Interspecific hybridization: progress and prospects. Proceedings of the workshop Africa/Asia joint research on interspecific hybridization between African and
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Asian rice species O. glaberrima and O. sativa. Mbe, Bouake (Côte d’Ivoire): WARDA/ADRAO. p 43-59. Kijima Y, Sserunkuuma D, Otsuka K. 2006. How revolutionary is the NERICA revolution? Evidence from Uganda. Dev. Econ. 44:252-267. Lafitte R, Blum A, Courtois G. 2003. Secondary traits to help identify drought-tolerant genotypes. In: Fischer KS, Lafitte R, Fukai S, Atlin Q. Hardy B, eds. Breeding rice for drought-prone environments. Los Baños (Philippines): International Rice Research Institute. p 37-48. Lamo J. 2010. Genetic studies on drought tolerance and grain shattering traits in rice. PhD thesis, University of KwaZulu-Natal. 205 p. Majerus V, Bertin P, Lutts S. 2007. Effects of iron toxicity on osmotic potential, osmolytes and polyamine concentrations in African rice Oryza glaberrima Steud. Plant Sci. 173:96-105. Singh BN, Mackill DJ. 1990. Genetics of leaf rolling under drought stress. In: Brar DS, ed. Proceedings of the 2nd International Rice Genetics Symposium. Manila (Philippines): International Rice Research Institute. p 159-166. Sirault XRR, Condon AG, Rebetzke GJ, Farquhar GH. 2007. Genetic analysis of leaf rolling in wheat. Proceedings of the 12th Wheat Breeding Society Assembly, Mildura, Australia. 

 

Acknowledgement
We gratefully acknowledge the Rockefeller Foundation for its financial assistance to the first author. 

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