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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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 V-
shaped 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. Breeding line Type Rice parent

002 CT16346-CA-20-M Interspecific Female
012 CT16344-CA-9-M Interspecific Female
Set D 072 CT16317-CA-4-M Interspecific Female
121 CK 73 O. sativa Female
147 IRAT104 O. sativa Male
197 IRAT257 O. sativa Male
129 CK73 O. sativa 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-20-
M/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 Parent 2 F1 F2 F1 on mid-parents F2 on F1

a
Var Mean Var b
Mean Mean Mean h2 h2

1 12 5.9 ± 1.5 121 3.2 ± 1.9 5.4 ± 0.9 4.5 ± 0.8 0.57 0.68
2 12 5.9 ± 1.5 129 4.0 ± 1.4 5.3 ± 1.0 4.7 ± 0.6 0.65 0.65
3 2 4.8 ± 1.3 121 3.2 ± 1.9 4.6 ± 1.2 4.6 ± 0.5 0.86 0.38
4 2 5.9 ± 1.1 129 4.0 ± 1.4 4.1 ± 1.1 4.4 ± 0.5 0.61 0.37
5 72 5.6 ± 1.1 147 3.9 ± 1.5 5.2 ± 1.2 4.6 ± 0.5 0.94 0.47
6 2 4.8 ± 1.3 197 3.6 ± 1.4 5.1 ± 0.8 4.6 ± 0.5 0.66 0.49
7 2 4.8 ± 1.3 147 3.9 ± 1.5 5.2 ± 1.1 4.8 ± 0.5 0.86 0.42
a
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. 
 
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Acknowledgement
We gratefully acknowledge the Rockefeller Foundation for its financial assistance to the first
author. 

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