Field Crops Research 99 (2006) 4858

www.elsevier.com/locate/fcr

Use of drought response index for identification of drought

tolerant genotypes in rainfed lowland rice
Makara Ouk a,b,*, J. Basnayake b, M. Tsubo b, S. Fukai b, K.S. Fischer b,
M. Cooper b,c, H. Nesbitt d
a

Cambodian Agricultural Research and Development Institute (CARDI), Phnom Penh, Cambodia
School of Land and Food Sciences, The University of Queensland, Brisbane, Qld 4072, Australia
c
Pioneer, DuPont Agriculture and Nutrition, 7250 N.W. 62nd Avenue, P.O. Box 552, Johnston, IA 50131-0552, USA
d
17 Heytesbury Road, Subiaco, WA 6008, Australia
b

Received 24 September 2005; received in revised form 13 March 2006; accepted 14 March 2006

Abstract
Drought is a major constraint for rice production in the rainfed lowlands in Southeast Asia and Eastern India. The breeding programs for
rainfed lowland rice in these regions focus on adaptation to a range of drought conditions. However, a method of selection of drought tolerant
genotypes has not been established and is considered to be one of the constraints faced by rice breeders. Drought response index (DRI) is based
on grain yield adjusted for variation in potential yield and flowering date, and has been used recently, but its consistency among drought
environments and hence its usefulness is not certain. In order to establish a selection method and subsequently to identify donor parents for
drought resistance breeding, a series of experiments with 15 contrasting genotypes was conducted under well-watered and managed drought
conditions at two sites for 5 years in Cambodia. Water level in the field was recorded and used to estimate the relative water level (WLREL)
around flowering as an index of the severity of water deficit at the time of flowering for each entry. This was used to determine if DRI or yield
reduction was due to drought tolerance or related to the amount of available water at flowering, i.e. drought escape.
Grain yield reduction due to drought ranged from 12 to 46%. The drought occurred mainly during the reproductive phase, while four
experiments had water stress from the early vegetative stage. There was significant variation for water availability around flowering among the
nine experiments and this was associated with variation in mean yield reduction. Genotypic variation in DRI was consistent among most
experiments, and genotypic mean DRI ranged from
0.54 to 0.47 (LSD 5% = 0.47). Genotypic variation in DRI was not related to WLREL
around flowering in the nine environments. It is concluded that selection for DRI under drought conditions would allow breeders to identify
donor lines with high drought tolerance as an important component of breeding better adapted varieties for the rainfed lowlands; two
genotypes were identified with high DRI and low yield reduction and were subsequently used in the breeding program in Cambodia.
# 2006 Elsevier B.V. All rights reserved.
Keywords: Rainfed lowland; Drought response index; Yield potential

1. Introduction
A large portion of the worlds poor farm in rainfed
systems where the water supply is unpredictable and
droughts are common. For example, in Thailand, about 50%
of all rice land is rainfed, and yield losses due to drought are
estimated to be between 11 and 58% (Jongdee et al., 1997).
* Corresponding author. Tel.: +855 23 219693; fax: +855 23 219800.
E-mail address: ou.makara@cardi.org.kh (M. Ouk).
0378-4290/$ see front matter # 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.fcr.2006.03.003

In Cambodia, drought can occur any time during the wet

season, and rice production may be reduced greatly, as
happened in 2004. In contrast to the irrigated rice system,
yield gains from crop improvement of rainfed rice have been
modest (Mackill et al., 1996), in part because there has been
little effort to breed and select for drought tolerance in these
target environments.
While many putative and specific traits for drought
tolerance such as osmotic regulation, root length, and root
penetration have been suggested for selection for drought

M. Ouk et al. / Field Crops Research 99 (2006) 4858

tolerance (Fukai and Cooper, 1995), there is yet no clear

evidence of their contribution to improved yield of rainfed
lowland rice under drought. Plant breeders, therefore, rely
on direct selection for grain yield (GY) in the target
environments as the main criterion for selection. However, a
genotype that produces high yield in a breeding experiment
exposed to drought conditions does not necessarily possess
traits for drought tolerance. The genotype may possess high
potential yield under water non-limiting conditions, or may
have escaped from drought by reaching maturity before
drought develops. Yield in drought-prone environments
may be considered to be affected by three components, viz.
yield potential, appropriate phenology and drought tolerance (Fig. 1). The contribution of drought tolerance to
actual grain yield may not be large, relative to that of
potential yield and phenology. For example, for chickpea in
the Mediterranean, the contribution of each component in 3
years varied from 37 to 69% for drought escape (appropriate
phenology), 147% for potential yield and 417% for
drought tolerance (Salim and Saxena, 1993). Thus, if
drought screening is based on grain yield, genotypes
selected may have high potential yield or appropriate
phenology, but not drought tolerance. The problem of
differences in crop phenology among genotpes may be
reduced by grouping genotypes into different maturity
groups, e.g. three groups in Cambodia, but there is still a 30
days variation within each maturity group (Ouk et al.,
2001).
A practical approach for selection of drought tolerance
parent is to use a measure or an index of the relative yield
of genotypes under stress to that under well-watered
conditions as an integrative measure of the complex of
traits that provide drought tolerance. For this purpose
Bidinger et al. (1987a,b) developed a drought response
index (DRI) to identify genotypes that are tolerant or
susceptible to drought and applied this to pearl millet
[Pennisetum americanum (L.) Leeke]. The DRI corrects
grain yield under drought for variation in flowering date

49

and potential yield under well-watered conditions, thus,

assuring that genotypes selected will have drought
tolerance traits. Drought response index has been used
in different crops such as rice, beans and chickpea (Abebe
et al., 1998; Salim and Saxena, 1993; Garrity and OToole,
1994; Yue et al., 2005). Pantuwan et al. (2002b) applied
this method for rainfed lowland rice in Thailand.
Genotypes differed in DRI, but the estimate of the DRI
was inconsistent across drought stress environments due to
differences in timing and intensity of water stress. If DRI
does not consistently identify tolerant genotypes, its use in
a breeding program will be limited. They used only four
stress environments, and the DRI performance needs to be
evaluated in a larger number of environments to determine
its consistency across environments.
The DRI for a given genotype may depend on the
available water at a critical development stage such as
flowering. Thus, a genotype may show higher DRI values
than others because they were exposed to a better water
environment in the experiments where genotypic performance was tested. If this is the case DRI is unlikely to be
consistent. Measuring the available soil water for the
different genotypes could help overcome this bias and thus
improve the consistency of the DRI. For example, if
genotypic variation in DRI is not due to water availability
among the genotypes, then genotypes with a high DRI would
indicate the possession of trait combinations contributing to
drought tolerance, and could be used as donors for
development of genotypes adapted to the target environments of the region.
Currently there is no information on the extent of
genotypic variation in drought tolerance among the rice
materials used in the breeding program in Cambodia. This
study aimed to: (1) determine whether genotypic differences
in DRI is related to water availability to the genotype
(escape) or to drought tolerance regardless of water
availability to the plants, and (2) identify genotypic variation
for DRI, of rainfed lowland rice genotypes and examine its
consistency across a wide range of water stress conditions at
different sites and over years in Cambodia.

2. Materials and methods

2.1. Field experiments

Fig. 1. Importance of genetic yield potential and flowering time in determining grain yield under various drought conditions.

2.1.1. Locations and treatments

The field experiments were conducted at the Cambodian
Agricultural Research and Development Institute (CARDI)
(latitude 118280 3600 N, longitude 1048480 2700 E) for 4 years
(19992002) and at Prey Veng province (latitude
118330 4000 N, longitude 1058320 4200 E) for 5 years (1998
2002) in Cambodia. Both sites are typical rainfed lowland
having Prateah Lang soil type (Plinthustalfs in USDA
classification and Acrisols or Luvisols or Planosols in FAO
classification) that covers 30% of the total rice growing areas

50

M. Ouk et al. / Field Crops Research 99 (2006) 4858

(White et al., 1997). The soil type has poor chemical and
physical attributes, and rice production is expected to be
lower than that on most other soils (White et al., 1997). The
experiment at each site-year combination consisted of two
water treatments; flooded, well-watered (WW) and water
stressed (WS) (drained fields to develop drought conditions),
which were located in adjacent paddies separated by a bund.
The WW treatment relied on rainfall, which was in some
experiments supplemented with irrigation to provide nonstress conditions. In general, the standing water depth in the
WW treatment fluctuated from 0 to 15 cm at Prey Veng and 0
to 20 cm at CARDI, although in some periods it was not
possible to maintain standing water at all times. In the WS
treatment, small canals (10 cm in depth) were dug
throughout the fields to collect water into a well 50 cm
deep, dug in the corner of the fields. The water was pumped
from the field during the drained period. The field was
drained from 1 to 3 weeks before 50% flowering of the
earliest maturity genotype to maturity for 19982000
experiments and from 2 weeks after transplanting to
maturity for 20012002 experiments to simulate a wide
range of drought conditions.
Fifteen genotypes (Table 1) were selected from a random
population consisting of 80 genotypes tested in an initial
trial at Prey Veng in 1998. The original 80 genotypes were
randomly sampled from Cambodian varieties and breeding
lines and Thai-ACIAR breeding lines; in 1999, 25 lines
were randomly sampled from the original 80 lines across
different groups of GY (low, intermediate and high) in
different maturity groups (early, intermediate and late
flowering); and in 2000, 15 lines were selected based on
different GY and different maturity groups from a set tested
in 1999. Of the 15 genotypes, seven were Cambodian
varieties, one was a Cambodian advanced breeding line, and
seven were Thai-ACIAR advanced breeding lines. Somaly
and Phka Rumchang are aromatic rice varieties. Three of the
genotypes were photoperiod insensitive (photoperiod

Table 1
Genotypes used in the experiments including the source and photoperiod
sensitivity index (PSI)
Gen. ID.

Genotype

Source

PSI

G1
G2
G3
G5
G8
G10
G13
G15
G16
G17
G18
G22
G24
G25
G26

IR46331-PMI-32-2-1-1
IR57514-PMI-5-B-1-2
IR66327-KKN-10-P1-3R-0
IR66327-KKN-25-P1-3R-0
IR66327-KKN-54-P1-3R-0
IR66327-KKN-8-P1-3R-0
IR66368-CPA-84-P1-3R-0
Bang Kuy (acc. 2865)
CAR4
CAR6
CIR158-B-B-SB-8-3-2
Khpor Daung
Somaly
Phka Rumchang
CAR3

Thai-ACIAR
Thai-ACIAR
Thai-ACIAR
Thai-ACIAR
Thai-ACIAR
Thai-ACIAR
Thai-ACIAR
Cambodia
Cambodia
Cambodia
Cambodia
Cambodia
Cambodia
Cambodia
Cambodia

0.45
0.24
0.55
0.24
0.53
0.48
0.76
0.71
0.73
0.73
0.30
0.81
0.77
0.77
0.73

sensitivity index (PSI) < 0.3, (Immark et al., 1997), four

were mildly photoperiod sensitive (0.3 < PSI < 0.7), and
the others were strongly photoperiod sensitive (PSI > 0.7).
When sowing date is delayed for 60 days from May, the
delay in flowering would be greater than 40 days in
photoperiod insensitive and less than 20 days in strongly
sensitive genotypes.
2.1.2. Experimental design
The experiments at each location and year were arranged
in three randomized complete blocks within the two water
treatments. For the analysis of variance, the water treatments
were treated as main-plots, with replicates within mainplots, and genotypes as sub-plots. The sub-plots were
1.2 m  3 m in size (six rows) with no extra row spacing
between them.
2.1.3. Crop management
The seeding date varied between early-June and midAugust, depending on the beginning of the wet season at the
location (Table 2). The beginning of the wet season is the
normal planting time in the target environments. Thirty days
old seedlings were transplanted by hand with three
seedlings per hill with 20 cm spacing between hills and
between rows. After 10 days, missing hills were again
transplanted. Irrigation was applied at transplanting to
maintain an above-ground water level of 35 cm in all
treatments and was continued until harvest in the WW
treatment and until the time of draining water in the WS
treatment.
The fields were fertilized with 603030 kg ha
1 of N
P2O5K2O. Nitrogen was applied as urea (46% N) on three
occasions (1/3 each at transplanting as a basal, at 30 days and
at 60 days after transplanting), while the P2O5 and K were
applied once as a basal application of triple super phosphate
(48% P) and of potassium chloride (60% K), respectively.
Weeds were controlled by hand three times in the WW and
four times in the WS treatments. The fourth weeding in the
WS treatment was 10 days after draining the water. No
pesticide was used.
2.1.4. Measurements
Days-to-flower (DTF) was measured as the time taken
from seeding to 50% of the 20 plants in the two center rows
of each plot to flower (anthers visible). At maturity, plants
from an area of 1.04 m2 were harvested at ground level from
the two center rows, leaving one hill at each end of the row as
a border. The harvested plants were sun-dried for several
days, and the grain threshed and then weighed. Sub-samples
of 150 g were taken and dried in an oven at 70 8C for 2 days.
The oven-dried samples were again reweighed to determine
the water content. Grain yield (GY) was calculated on a dryweight basis.
The level of the paddy water table above or below the soil
surface was recorded weekly using PVC tubes placed in each
corner of the main water treatments plots.

0.75
0.56
0.68

2002

WW
WS
13-Aug
2.15
1.47
32
Moderate
From vegetative stage

51

The level of the paddy water table above or below the soil
surface was recorded weekly using PVC tubes placed in each
corner of the main water treatments plots.

0.62
0.60

0.68

0.95

0.35

0.73

2001

WW
WS
4-Jun
2.63
1.47
44
Severe
From vegetative stage

2000

WW
WS
31-Jul
2.42
2.13
12
Mild
Short grain
filling stage
WW
WS
2-Jun
2.72
2.07
24
Mild
From booting
stage

0.62
0.73
0.84
0.42
0.32

0.48

Yact;i
Yest;i
S:E: of Yest

(1)

where Yact is the actual grain yield under the WS treatment

for each genotype, Yest is the estimated GY for each genotype under the WS treatment, and S.E. of Yest is the standard
error of estimated GY of all genotypes. Estimated GY of a
specific genotype i, Yest,i, was derived from a multiple
regression analysis as follows:
(a) when the relationship between GY in the WS treatment
(Yact) and DTF in the WW treatment (FL) is not
quadratic
Yest;i a bY p;i cFLi
(2a)
(b) when the relationship between GY in the WS treatment
(Yact) and DTF in the WW treatment (FL) is quadratic
Yest;i a bY p;I cFLi dFL2i

(2b)

where Yp,i is the potential GY of genotype i, as measured

under the WW treatment, and FLi is the DTF of genotype i under the WW treatment, and a, b, c and d are
regression parameters estimated by least square methods. Examples of such relationships are shown in Fig. 2.

0.84

0.93

0.88

0.89

0.71

0.65

0.85

5-Jul

0.76
0.80
0.11
0.85
0.82
(b) Genotypic variation
Heritability
Phenotypic correlation

1.98

(a) Drought condition

Sowing date
GY (t ha
1)
YR (%)
Drought severity
Period of drought

WW
WS
11-Jul
1.93
1.52
21
Mild
From grain
filling stage

WW

1.08
46
Severe
From booting
stage

WW
WS
4-Jun
2.21
1.77
20
Mild
From vegetative stage
WW
WS
4-Jun
2.46
1.71
30
Moderate
From vegetative stage
WW
WS
18-Jul
1.49
1.17
21
Mild
From flowering stage
WS

2002
2001
1999
Prey Veng (PV)

1998

2.2.1. Drought response index (DRI)

The DRI for a specific genotype i (i = 1, . . ., g), DRIi was
calculated using a modified equation of Bidinger et al.
(1987b) as follows:
DRIi

2000

CARDI (CA)

1999

2.2. Definitions and calculations

Location year

Table 2
Mean grain yield (GY) at Prey Veng (PV) and CARDI (CA) under the well-watered (WW) and water-stress (WS) treatments, broad sense heritability estimated on a line mean basis, relative grain yield reduction
(YR), phenotypic correlation between WW and WS experiments, severity of drought and period of drought for the 15 rice genotypes shown in Table 1

M. Ouk et al. / Field Crops Research 99 (2006) 4858

Bidinger et al. (1987b) computed a DRI for a genotype

from the mean GY of the genotype across replicates in the
drought trial and the mean GY and DTF of the genotype in
the well-watered trial. In this study, a mean value for GY and
DTF of the genotype in the WW treatment was computed
and used to calculate the DRI separately for each replicate in
the WS treatment. The resulting experimental error for the
DRI is a combined error of the genotype mean error from the
WW treatment and the single replicate error from the WS
treatment. An advantage of this method, over the approach
used by Bidinger et al. (1987b), for computing the DRI for
individual replicates in the WS treatment is that with
replicate measurements in the WS treatment, the DRI could
be more fully analysed by ANOVA to study the consistency
of the DRI values in different environments.
2.2.2. Relative water level (WLREL)
The relative water level (WLREL) was determined as the
difference in paddy water level (WL) between the WW and
WS treatments and used as an index of water-deficit to
explain the yield reduction (YR) due to drought. The WLREL

52

M. Ouk et al. / Field Crops Research 99 (2006) 4858

(d) when WL of both water conditions was below the soil

surface, the difference in level between the two water
treatments was taken as WLREL. Thus
WLREL WLWS
WLWW

(3d)

when WLWW < 0; WLWS < 0

Since WL was measured weekly, WL between measurement days was estimated by interpolation from the two
consecutive measurements. WLREL was averaged over 6
weeks around flowering (3 weeks before and after flowering
date) of a genotype or around mean flowering date of an
experiment as water deficit during this 6 week period was
found to best correlate with yield reduction due to water
stress.
2.3. Data analysis

Fig. 2. The relationship between grain yield and flowering date at Prey
Veng in 2000 and 2002. Three symbols indicate introduction from ThaiACIAR (circle), Cambodia varieties (square) and Cambodian advanced
lines (triangle). Individual replicate values are shown.

was calculated based on four water conditions of the WW

and WS treatments, as follows:
(a) when WL of both water treatments was above the soil
surface, there was no water deficit, and hence the two
water treatments had the same water availability. Thus
WLREL 0

when WLWW  0; WLWS  0

(3a)

where WLWW and WLWS are free water level for the
WW and WS treatments.
(b) The case of WLWW  0 with WLWS < 0 occurred
frequently after water was drained in the WS field.
WLREL for these water conditions was calculated as
WLREL WLWS when WLWW  0; WLWS < 0 (3b)
(c) The case of WLWW < 0 with WLWS  0 was rare; the
occurrence was only a few days during the flowering
periods in a few environments. The effect on the
determination of WLREL was, therefore, minimal.
WLREL for these water conditions was calculated as
WLREL
WLWW

when WLWW < 0; WLWS  0

(3c)

An analysis of variance (ANOVA) was conducted on the

data of DTF, GY and DRI using a residual maximum
likelihood (REML) program (Robinson et al., 1982). It was
assumed that the genotypes represented a random sample of
rainfed lowland rice genotypes and that the sites and years
were a random sample of those used by the Cambodian
breeding program. The water treatment was a fixed variable.
The treatment means for all random variables were
computed as best linear unbiased predictors (BLUPs).
REML software (Robinson et al., 1982) was used to
calculate the BLUPs. Two main analyses were performed for
DTF and GY. The first was a combined analysis over both
water treatments to detect the effects of water, genotype and
their interaction effects at each site. In the second analysis, a
combined analysis over sites or years was performed to
detect the effects of site or year and genotype and their
interaction effects for DTF, GY and DRI.

3. Results
3.1. The water environments and types of drought
In 1998 at Prey Veng (PV98), rainfall was well distributed
until late December (Fig. 3). There was standing water in the
WW treatment until maturity, whereas there was no standing
water in the WS treatment from early November. At both
sites (CARDI and Prey Veng), rain ceased by early
November in 1999 and 2001, while it continued until
December in 2000 and 2002. In 1999, there was no standing
water in the WS treatment soon after draining in late
September at both sites. In the WS treatment in 2000, there
was no standing water from early November and WL
decreased during the flowering period at both sites. In the
WS treatment in 2001, there was no standing water
immediately after draining in early July and water level
was low in Prey Veng. In CA01, the crop was completely

M. Ouk et al. / Field Crops Research 99 (2006) 4858

53

Fig. 3. Rainfall (bars) and water levels under the well-watered (WW) (*) and the water-stress (WS) (*) treatments at CARDI and Prey Veng from 1998 to
2002, Cambodia.

submerged by a flash flood from 13 to 16 October with

maximum water depth of 95 cm. This period coincided with
the 1 week of mean flowering date for the early maturity
group (7 October for the WW treatment and 11 October for
the WS treatment) and affected those genotypes that

flowered at that time. In the WS treatment in 2002, there

was no standing water from early October onwards. The
water was closer to the surface at Prey Veng.
The draining technique was successful in providing
various drought conditions, which ranged from drought after

54

M. Ouk et al. / Field Crops Research 99 (2006) 4858

booting or during the grain filling in 19982000 to

prolonged drought from vegetative stage in 20012002
(Table 2). The yield reduction (YR) varied from 12% in
CA00 up to 46% in PV99 and is used as a measure of the
severity of the imposed droughts. Thus, the variation in the
severity of the imposed droughts ranged from mild drought
for YR  24%, moderate drought for YR between 30 and
32%, and severe drought for YR  44%. The WLREL around
mean flowering date of WS treatment varied from
6 to

7 cm in PV02 and CA00 to
28 to
36 cm in PV99 and
CA99. Except for CA99, YR was related to WLREL
(YR = 6.72
1.15 WLREL, r = 0.59*, n = 8).
In CA99, although WLREL was low, water appeared to be
available to the plants in the WS treatment. The amount of
the 3-month rainfall was 2993% higher in CA99 than in the
other locations. This high rainfall during growth may have
resulted in greater lateral water movement in the paddy
fields in CA99 than in other experiments. The lateral water
inflow may have increased soil wetness of the topsoil in the
WS field during the flowering period (mid October to mid
November), although the free water level was low. After
1999 a new site was selected for CARDI WS treatment
where lateral water flow would be small.
3.2. Genotypic variation and heritability (broad sense)
estimates for GY, DTF and DRI
Broad sense heritability (expressed on a line-mean basis)
for GY was estimated for nine pairs of WW and WS
treatments (Table 2). In the WW treatment, the heritability
for GY ranged from moderate 0.42 in PV02 to 0.95 in CA01.
In the WS treatment, the line mean heritability ranged from
0.11 in PV98 to 0.89 in PV00. The heritability of PV98-WS
treatment was low because of the high error variance due to
weed competition with some lines in one replication. The
mean GY of the 15 genotypes grown under the WS and WW
treatments were correlated (r > 0.51 for P < 0.05) in most
experiments (Table 2). Associations were stronger in mild
drought (r > 0.80 in PV98, PV00, CA99) and low-medium

in moderate and severe drought (r < 0.8 in PV99, PV01,

CA01, CA02). However, the association was intermediate
(CA00) to negative (PV02) despite the drought was
classified as mild in these experiments. In PV02, the
drought occurred at the early vegetative stage and the early
flowering, high yield potential genotypes were affected by
the drought whereas the late flowering, lower potential
yielding genotypes were not affected. Thus, there was a
negative relationship between GY under the WW and WS
treatments in PV02. However, mean genotype grain yield
across nine experiments was highly correlated between the
WW and WS treatments (r = 0.73**), indicating the
importance of potential yield for genotypic performance
under drought conditions.
Genotypes varied for GY-WS, GY-WW and DTF and
DRI (Table 3). The main variations of year (Y), site (S), and
interaction variations of Y  S, G  Y and G  S were not
significant for DRI, whereas Y  S, G  S and G  Y  S
were significant for GY-WS and GY-WW. The three-way
interaction (G  Y  S) was significant and had the highest
variance component among all interaction terms that
involved genotype for DRI, GY-WS, GY-WW and DTF
(Table 3). The ratio of the variance components s 2GE =s 2g
was 0.9 for GY-WW, whereas the ratio was 2.7 for GY-WS
and 4.4 for DRI. DTF under WS condition had the smallest
s 2GE =s 2g ratio (0.35).
The G  W  S  Y component of variance for GY was
about 50% higher than that for the genotype component and
large relative to the other interaction variance components.
The ratio of the variance components s 2GE =s 2g was 2.5.
Thus, the s 2GE =s 2g ratio for combined treatments of WW and
WS was similar to that obtained under WS and greater than
that obtained under WW. Broad sense heritability (on linemean basis) for GY-WW and GY-WS treatments, DTF under
the WS treatment and DRI were estimated from the
combined analyses (Table 3).
DRI varied with genotype in seven of the nine
experiments (Table 4). The mean DRI across the experiments also differed among the 15 genotypes. The mean DRI

Table 3
Components of variance  standard errors (s2  S.E.), and the ratio of genotype-by-environment (year and site) interaction and genotype component of
variance (s 2GE =s 2g )a for DRI and grain yield of WS and WW conditions for 15 genotypes grown under two water treatments at two sites (CARDI and Prey Veng)
and DTF for WS treatment in 5 years (19982002) in Cambodia
GY-WS

S.E.

GY-WW

S.E.

DTF-WS

S.E.

DRI

S.E.

Year (Y)
Site (S)
YS
Genotype (G)
GY
GS
GSY
Residual

0.00
0.01
0.24
0.02
0.00
0.01
0.05
0.09

0.14
0.09
0.19
0.01
0.01
0.01
0.02
0.01

0.01
0.09
0.09
0.10
0.02
0.00
0.07
0.13

0.06
0.16
0.08
0.05
0.02
0.01
0.02
0.01

121
0
220
86
0
1.28
29.15
6.89

200
83
177
34
4.83
3.5
6.63
0.62

0.00
0.00
0.00
0.06
0.00
0.00
0.25
0.59

0.02
0.01
0.02
0.03
0.07
0.04
0.10
0.05

G  Ea/G
Heritability

2.69
0.64

0.90
0.86

0.35
0.95

4.41
0.56

a
The G  E interaction component of variance was estimated as the sum of the genotype-by-year, genotype-by-site and genotype-by-year-by-site
components for the analysis of two sites in 5 years.

M. Ouk et al. / Field Crops Research 99 (2006) 4858

55

Table 4
The drought response index (DRI), LSD 5% values for mean comparison, broad sense heritability (h2) for DRI, genotype-by-environment/genotype ratio of the
estimated components of variance (s 2GE =s 2g ), when 15 genotypes were grown at CARDI (CA) for 4 years and at Prey Veng (PV) for 5 years

Mean yield reduction% (YR%) and the mean relative water level (WLRWL) for each genotype are also shown on the last two columns. Shading and bold indicate
the values smaller than
1.00 and higher than 1.00. ns = not significant.

varied from 0.47 (CAR3) and 0.43 (IR57514-PMI-5-B-1-2)

to
0.54 (IR46331-PMI-32-2-1-1) and
0.50 (CAR4). The
effects of the genotype by environment interaction for DRI
were significant (P < 0.01). The broad sense heritability of
DRI in the different environments varied from 0.23 at PV98
to 0.75 at CA02.
The WLREL for the 15 genotypes averaged over the nine
environments, ranged between
16 and
25 cm (Table 4).
The variety CAR3 had high DRI (0.47) and 18% yield
reduction (YR) associated with a WLREL of
24 cm. On the
other hand CAR4 had a low DRI (
0.50) and 35% YR with
the same WLREL (
24 cm). These two genotypes showed
contrasting levels of DRI and thus drought tolerance under
similar water levels with a small variation in phenology.
Similarly IR57514-PMI-5-B-1-2 and IR46331-PMI-32-2-11 differed in DRI but the genotypes had a similar water
supply and similar phenology. These results indicate that
genotypic variation in DRI was not related to water
availability and DRI is a measure of drought tolerance.
Mean grain yield of each genotype was not strongly
related to mean flowering date in either well-watered or
water stressed conditions (Fig. 4). Three early maturing

genotypes from Thailand had high grain yield particularly

under well-watered conditions. There was a positive
association between mean grain yield under the WS
treatments and the mean DRI of genotypes (r = 0.59*,
n = 15) across the nine environments. When each experiment was considered, the correlation was positive in all
experiments, but some were not significant (Table 5). On the
other hand, the genotypic variation for DRI was not related
to WLREL or flowering date.

4. Discussion
Although the types of drought varied in timing with crop
development, from prolonged drought from the vegetative
stage to a short duration in the grain filling stage, and their
effects on grain yield, the actual stress mainly developed late
in the season. The drought effect ranged from a 12 to 46%
reduction in GY. In similar environments in Thailand,
Wonprasaid et al. (1996) obtained a 40% reduction in GY
using the method of draining the paddies about 1 month
before flowering, while Pantuwan et al. (2002a) obtained

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M. Ouk et al. / Field Crops Research 99 (2006) 4858

Fig. 4. Relationship between mean grain yield of each genotype and mean
days to flower under well-watered and drained (water stressed) conditions.
Genotypes are grouped into Thai-ACIAR (circle), Cambodian varieties
(square) and Cambodian advanced lines (triangle).

reductions from 19 to 80%. In other studies in Cambodia,

Basnayake et al. (2004) estimated yield reductions due to
drought from 9 to 51% for three maturity groups of
genotypes in multi-location trials conducted in 3 years
(20002002) in the target environments. Thus, the ranges on
drought intensity established in the trials in the current study
are representative of the target population of environments.
The controlled drought treatments were used to estimate
the DRI of a diverse range of genotypes grown in the rainfed
lowlands in Cambodia. There was a significant difference
among the genotypes for DRI, and they were reasonably

consistent across different drought environments, indicating

this technique can be used to identify genotypes that confer
drought tolerance. This finding differs from that of Pantuwan
et al. (2002b) who suggested that there was no consistency of
DRI across varying drought environments. In their experiments, drought developed rather quickly in some experiments where the plants had developed a large biomass by the
time of the drought, resulting in a sharp reduction in yield,
particularly in later maturing genotypes. In the present
experiments, drought developed rather slowly in all
experiments, and hence the phenology effect on grain yield
reduction was small to insignificant.
Genotypes with high DRI values were identified. One
genotype (IR57514-PMI-5-B-1-2), which had the second
highest DRI value in the study here, was also found by
Jearakongman et al. (1995) to be drought tolerant. This
genotype also performed well under diverse rainfed lowland
environments in northeast Thailand (Romyen et al., 1998)
and in Lao PDR (Inthapanya et al., 2001). This result
suggests that it is possible to use IR57514-PMI-5-B-1-2 as a
parent in a rainfed lowland breeding program for the
development of drought tolerant varieties. This genotype is
one of the donors used in the Thai rice-breeding program for
drought tolerance. The measurement of soil water level
indicates that variation in DRI was not associated with water
availability. Thus, DRI appears useful for identifying
drought tolerant genotypes. In addition, because DRI was
not confounded by GY or DTF under the WW treatment,
selection for high DRI should not have negative effects on
yield potential and maturity.
Determination of DRI requires an irrigated control trial. It
is also costly, and hence the technique needs to be used
strategically in the breeding program. The use of DRI is
likely to be most beneficial in selecting parents for
development of drought resistant populations, particularly
when flowering time and/or yield potential vary greatly
among the tested genotypes. Even if genotypes are grouped
according to maturity, 30 days difference within a group
would have large effect on grain yield under drought, and
DRI can be used to correct for the difference in phenology. If
variation in phenology and potential yield is small, grain
yield can be used directly as a selection criterion. Also if
drought develops rather quickly and severely, then DRI is
often associated directly with yield, and hence there is no
need to use DRI.

Table 5
Phenotypic correlation between grain yield and flowering date under stress conditions and DRI of 15 genotypes at Prey Veng (19882002) and CARDI (1999
2002)
Grain yield (t ha
1)
Location

Prey Veng

Year

1998

1999

2000

2001

2002

1999

CARDI
2000

2001

2002

Days to flower
DRI

0.31
0.55

0.36
0.49

0.23
0.39

0.61
0.67

0.52
0.49

0.05
0.84

0.04
0.56

0.23
0.51

0.11
0.60

Correlation is significant when r > 0.49.

M. Ouk et al. / Field Crops Research 99 (2006) 4858

This study developed a protocol for estimating DRI that

can be applied routinely in the Cambodian breeding
program. Whereas the method proposed by Bidinger et al.
(1987b) uses threshold values for the upper and lower 10%
of the normal distribution (Z = +1.3 and
1.3) to identify
drought tolerant (DRI > 1.3) and susceptible (DRI <
1.3)
genotypes, the method described in the present study
estimates DRI for each replication and hence allows a
comparison of all of the genotypes based on significant
variation for DRI among genotypes. Researchers can use
this test to judge the level of consistency of DRI among the
genotypes growing under different environments and
identify genotypes with high and consistent DRI values
for use as parents in a crossing program.
This study did not attempt to understand the underlying
mechanisms and traits that contribute to the variation in DRI.
Jearakongman et al. (1995) found that the high DRI
genotype IR57514-PMI-5-B-1-2 had greater green leaf
retention under the prolonged drought. The value of DRI as a
measure of drought tolerance will be enhanced once the
mechanisms and traits for the variation in DRI are better
understood. However, for now, plant breeders can screen and
use DRI along with yield potential and appropriate maturity
to identify parents to improve the performance of rainfed
lowland rice under variable moisture environments. This
work identified CAR3 as drought tolerant and CAR4 as
drought susceptible among Cambodian varieties. CAR4 has
low DRI and high yield reduction under poor water
conditions. Studying the performance of drought adaptation
of two donor parents would assist to quantify the importance
of DRI as a trait for parent selection.

5. Conclusions
The establishment of managed drought conditions by
draining the rainfed lowland paddies allows breeders to
select drought tolerant genotypes. Genotypes with high DRI
and high GY performed consistently across the range of
water environments sampled in the experiments.. While soil
water level at around flowering explained a large variation in
grain yield under various drought environments, it was not
related to genotypic variation in DRI. Thus, DRI was shown
to be associated with drought tolerance. When these
genotypes with high DRI are used as parents, they are
likely to contribute to development of varieties adapted to
the target drought-prone environments.

References
Abebe, A., Brick, M.A., Kirkby, R.A., 1998. Comparison of selection
indices to identify productive dry bean lines under diverse environmental conditions. Field Crops Res. 58, 1523.
Basnayake, J., Ouk, M., Thun, V., Kang, S., Pith, K.H., Fukai, S., Men, S.,
Fischer, K., 2004. Measurement and management of genotype-environment interaction (G  E) for the improvement of rainfed lowland rice

57

yield in Cambodia. In: Proceedings for the 4th International Crop

Science Congress, Brisbane, Australia, 26 September1 October, p.
239., http://www.cropscience.org.au.
Bidinger, F.R., Mahalakshmei, V., Rao, G.D.P., 1987a. Assesment of
drought resistance in pearl millet [Pennisetum americanum (L.)
Leeke]. I. Factors affecting yield under stress. Aust. J. Agric. Res.
38, 3748.
Bidinger, F.R., Mahalakshmei, V., Rao, G.D.P., 1987b. Assessment of
drought resistance in pearl millet [Pennisetum americanum (L.) Leeke].
II. Estimation of genotypes response to stress. Aust. J. Agric. Res. 38,
4959.
Fukai, S., Cooper, M., 1995. Development of drought-resistant cultivars
using physio-morphological traits in rice. Field Crops Res. 40, 67
86.
Garrity, D.P., OToole, J.C., 1994. Screening rice for drought resistance at
the reproductive phase. Field Crops Res. 39, 99110.
Immark, S., Mitchell, J.H., Jongdee, B., Boonwite, C., Somrith, B., Polvatana, A., Fukai, S., 1997. Determination of phenology development in
rainfed lowland rice in Thailand and Lao PDR. In: Fukai, S., Cooper,
M., Salisbury, J. (Eds.), Breeding Strategies for Rainfed Lowland Rice
in Drought-prone Environments. Proceedings of an International Workshop held at Ubon Ratchathani, Thailand, 58 November 1996. ACIAR
Proceedings, vol. 77, pp. 8996.
Inthapanya, P., Sipaseuth, Sihavong, P., Sihathep, V., Chanphengsay, M.,
Fukai, S., Basnayake, J., 2001. Genotipic performance of rainfed
lowland rice under different fertilizer conditions in laos. In: Fukai,
S., Basnayake, J. (Eds.), Increased Rainfed Lowland Rice Production
in the Mekong Region. Proceedings of an International Workshop held
at Vientiane, Laos, 30 October2 November 2000. ACIAR Proceedings 101, pp. 191200.
Jearakongman, S., Rajatasereekul, S., Naklang, K., Romyen, P., Fukai, S.,
Skulkhu, E., Jumpaket, B., Nthabutr, K., 1995. Growth and grain yield
of contrasting rice cultivars grown under different conditions of water
availability. Field Crops Res. 44, 139150.
Jongdee, S., Mitchell, H.J., Fukai, S., 1997. Moddleing approach for
estimation of rice yield reduction due to drought in Thailand. In:
Fukai, S., Cooper, M., Salisbury, J. (Eds.), Breeding Strategies for
Rainfed Lowland Rice in Drought-prone Environments. Proceedings of an International Workshop held at Ubon Ratchathani,
Thailand, 58 November, 1996. ACIAR Proceedings, vol. 77,
pp. 6573.
Mackill, D.J., Coffman, W.R., Garrity, D.P., 1996. Rainfed lowland rice
improvement. IRRI, PO Box 933, Manila, Philippines, pp. 242.
Ouk, M., Men, S., Nesbitt, J.H., 2001. Rice production systems in Cambodia. In: Fukai, S., Basnayake, J. (Eds.), Increased Rainfed Lowland
Rice Production in the Mekong Region. Proceedings of an International
Workshop, Vientiane, Loas, 30 October2 November, 2000. ACIAR
Proceedings, 101, pp. 4352.
Pantuwan, G., Fukai, S., Cooper, M., Rajatasereekul, S., OToole, J.C.,
2002a. Yield response of rice (Oryza sativa L.) genotypes to different
types of drought under rainfed lowlands. 1. Grain yield and yield
components. Field Crops Res. 73, 153168.
Pantuwan, G., Fukai, S., Cooper, M., Rajatasereekul, S., OToole, J.C.,
2002b. Yield response of rice (Oryza sativa L.) genotypes to different
types of drought under rainfed lowlands. 2. Selection of drought
resistant genotypes. Field Crops Res. 73, 169180.
Robinson, D.L., Thompson, R., Digby, P.G.N., 1982. REML, a program for
the analysis of non-orthogonal data by restricted maximum likelihood.
In: COMPSTAT Part II (supplement), Physica Verlag, Vienna, pp. 231
232.
Romyen, P., Hanviriyapant, P., Rajatasereekul, S., Khunthasuvan, S., Fukai,
S., Basnayake, J., Skulkhu, E., 1998. Lowland rice improvement in
northern and northeast Thailand. 2. Cultivar differences. Field Crops
Res. 59, 109119.
Salim, S.N., Saxena, M.C., 1993. Adaptation of spring-sown chickpea to the
Mediterranean basin. 2. Factors influencing yield under drought. Field
Crops Res. 34, 137146.

58

M. Ouk et al. / Field Crops Research 99 (2006) 4858

White, F.P., Oberthur, T., Pheav, S., 1997. Soils and Rice. In: Nesbitt, H.J.
(Ed.), Rice Production in Cambodia. International Rice Research
Institute (IRRI), Manila (Philippines), pp. 2130.
Wonprasaid, S., Khunthasuvon, S., Sittisuang, P., Fukai, S., 1996. Performance of contrasting rice cultivars selected for rainfed lowland condi-

tions in relation to soil fertility and water availability. Field Crops Res.
47, 267275.
Yue, B., Xiong, L., Xue, W., Xing, Y., Luo, Lijun, Xu, C., 2005. Genetic
analysis for drought resistance of rice at reproductive stage in field with
different types of soil. Theor. Appl. Genet. 111, 11271136.