Field Crops Research 162 (2014) 3038

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Field Crops Research

journal homepage: www.elsevier.com/locate/fcr

Biomass accumulation and partitioning of newly developed Green

Super Rice (GSR) cultivars under drought stress during the
reproductive stage
Manuel Marcaida III a , Tao Li a, , Olivyn Angeles a , Gio Karlo Evangelista a ,
Marfel Angelo Fontanilla a , Jianlong Xu b , Yongming Gao b , Zhikang Li b , Jauhar Ali a,
a
b

International Rice Research Institute, DAPO 7777, Metro Manila, Philippines

Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, 12 South Zhong-Guan-Cun St., Beijing 100081, China

a r t i c l e

i n f o

Article history:
Received 17 January 2014
Received in revised form 19 March 2014
Accepted 20 March 2014
Available online 12 April 2014
Keywords:
Green Super Rice
Introgression breeding
Drought tolerance mechanism
Biomass and yield advantage

a b s t r a c t
Drought is a major abiotic threat in rice production; thus, there is a need to develop adaptable rice varieties that can withstand drought stress and still produce high yield in non-stressed environments. Green
Super Rice (GSR) cultivars address this issue. These cultivars are bred through an innovative introgression
breeding strategy that requires less irrigation water and chemical inputs without compromising grain
quality and yield. This study veried the physiological efciency and performance of newly developed GSR
cultivars that previously showed favorable response to drought during advanced yield trials. Five droughttolerant GSR cultivars and two checks were subjected to continuously ooded (CF) and drought-stressed
environments during the dry seasons of 2011 and 2012 at the International Rice Research Institute (IRRI)

experimental farms in Los Banos,

Philippines. The cultivars ability to allocate assimilates and accumulate biomass under drought stress during the reproductive stage was veried. Leaf area index (LAI),
biomass dry weight, and panicle yield were measured at the panicle initiation (PI), owering (FL), and
physiological maturity (PM) of the sample cultivars. All the cultivars performed satisfactorily in the CF
environment with grain yield ranging from 5 to 11.5 tons ha1 . Water stress during the reproductive stage
signicantly reduced grain yield by 7588% in the moderate drought (soil water tension between 100 and
300 kPa in upper 15 cm soil layer) and 7796% in the severe drought (soil water tension >300 kPa in upper
15 cm soil layer) experiments. The shortened reproductive duration mainly contributed to the signicant
reduction in yield under drought stress. Two GSR cultivars, GSR IR1-5-S10-D1-D1 and IR83142-B-19-B,
responded well in severe drought environments, with grain yield almost similar to the drought check
(1.79 tons ha1 ). Under moderate drought stress, there was a relative yield advantage of 25% and 40%
for the two GSR cultivars over the drought check, respectively. Yield advantage across environments,
varying from fully irrigated to drought-stressed, was 3136%. These two GSR cultivars were effective in
mobilizing stored carbohydrates from the vegetative organs to the panicles and not shortening the duration from owering to maturity, to allow all reserved carbohydrates be allocated to storage organs as a
mechanism to cope with drought stress. Lower leaf area index (LAI), which allowed balanced biomass
accumulation and lower transpiration, without a signicant decrease in grain lling duration, was another
drought-coping strategy. These physiological responses and characteristics apparently enabled the GSR
cultivars to withstand drought stress; these are key indicators for varietal selection in drought-prone
environments, particularly in severe drought stress in the reproductive stage. Despite the poor ability of
some cultivars to cope with severe drought, three out of ve selected GSR cultivars produced grain yield
(2.02.9 tons ha1 ) that was the same or higher than the drought check in moderate drought stress. The
introgression breeding technique applied in the newly developed drought-tolerant cultivars through the
GSR breeding strategy was found to be effective. It could produce high yields in both CF and water-limited
environments, and thus, it could serve as a model for other breeding programs to adopt.
2014 Elsevier B.V. All rights reserved.

Corresponding authors. +63 25805600.

E-mail addresses: T.Li@irri.org (T. Li), J.Ali@irri.org (J. Ali).
http://dx.doi.org/10.1016/j.fcr.2014.03.013
0378-4290/ 2014 Elsevier B.V. All rights reserved.

M. Marcaida III et al. / Field Crops Research 162 (2014) 3038

1. Introduction
Ninety percent of rice is produced and consumed in Asia
(Timmer, 2010; Rejesus et al., 2012) where it is the staple food
of more than 4 billion people. Sustaining a stable rice supply in
the coming years is going to be a challenging task with decreasing
sources of water, arable land, and fertilizer input. Furthermore, climate change associated with an increasing frequency of extreme
climate events such as drought, oods, and cold or heat waves
adds up another dimension in the existing challenge (Easterling
et al., 2007; Meehl et al., 2007; USGCRP, 2009; CCSP, 2008). Rice
needs to be produced stably without creating production uctuations as experienced during the global food shortage in 2007 and
2008 (UNCTAD, 2008; FAO, 2010). Among the effects of climate
change, drought is a serious threat to food security, especially
in rainfed lowlands where 75% of the global rice supply is produced (Maclean et al., 2002). According to Pandey and Bhandari
(2009), 20% of rice-producing areas in Asia are affected by drought.
Adoption of high-yielding varieties in rainfed areas is constrained
because farmers are hesitant to take the risk of not applying
irrigation (Latte et al., 2007). Also, most of the drought-resistant
varieties that have been developed have a lower yield potential
than high-yielding rice varieties already adopted by the farmers
(Latte et al., 2007). Among the rice production constraints that
need immediate attention is the need to develop drought-tolerant
varieties suitable for rainfed and irrigated lowlands where drought
is going to recur frequently in the coming years (Latte et al.,
2007). It is known that drought occurrence is cyclic in nature and
varieties need to perform better in both normal and drought conditions. Currently, we nd very few rice varieties available for such
conditions tha have been developed through conventional breeding approaches. Moreover, conventional breeding requires more
time to develop high-yielding, drought-tolerant varieties because
of trait complexities. Among those are the challenges in identifying target environments, the interaction of drought tolerance with
environments, and the availability of the appropriate screening
methodology (Latte et al., 2006). An alternative approach has to be
in place to develop new varieties that could cope with the urgency
to produce favorable yield for farmers, whether in drought stress
or in fully irrigated conditions. One such method is introgression
breeding as adopted for developing these drought-tolerant materials.
Introgression breeding efforts to identify promising lines that
could survive drought, salinity, and submergence were initiated
in 1998 at the International Rice Research Institute (IRRI) by
Dr. Zhikang Li under the International Rice Molecular Breeding
Program (Li et al., 2005; Ali et al., 2006; Latte et al., 2006). However, the work on drought tolerance using an IR64 recipient was
advanced to design QTL pyramiding and to identify several pyramiding lines (PDLs) that could withstand severe drought conditions
(Guan et al., 2010). Furthermore, research on developing varieties
that could tolerate multiple stresses was initiated in 2008 through
the Green Super Rice (GSR) breeding strategy. It is an innovative
breeding approach that addresses the challenge of making poor
farmers benet from high-yielding varieties with limited resources
(Ali et al., 2013).
The Green Super Rice (GSR) breeding program at IRRI has
been successful in developing drought-tolerant cultivars (Ali et al.,
2012) with yields that are comparable with harvests in favorable environmental conditions. The new breeding strategy has
developed several multiple abiotic stress-tolerant cultivars that
are currently tested under the National Cooperative Testing Trials in the Philippines. Newly bred GSR lines were developed by
screening BC1 F2 bulk populations with Huanghuazhan (HHZ) background. They were introgressed from 27 donors to varied abiotic
stress conditions, including severe drought over three rounds of

31

intensive selections, before being nominated to preliminary (two

seasons) advanced yield trials (two seasons). These lines were
then subjected to three stress conditionsdrought, irrigated, and
low-nutrient input. Drought lines selected through this innovative
breeding strategy have already shown their relative performance.
More than 87 IRRI-bred GSR lines and 112 CAAS (Chinese
Academy of Agricultural Sciences)-bred GSR materials have been
distributed to national research partners in 16 countries in Asia
and Africa. The GSR materials are also being provided globally
through IRRIs International Network for Genetic Evaluation of Rice
(INGER), with trials on a regular basis, since 2009. Field trials with
GSR cultivars, both in IRRI and Asia, have already demonstrated
good performance in both irrigated and drought conditions (Ali
et al., 2012). Development and adoption of high-yielding droughttolerant cultivars that can withstand severe drought stress and
still produce higher yields in normal conditions (without drought)
appear to be a lasting solution in rainfed conditions. There is, then,
an urgent need to test the physiological efciency of these cultivars
and their performances under drought and irrigated conditions.
The improvement of harvest index and increase in dry matter
accumulation were reported as two main factors that enhance rice
yield under drought and irrigated conditions (Guan et al., 2010).
Moreover, high dry matter accumulation in the reproductive stage
also proved to greatly correlate to high yield under drought. The
study of Wu et al. (2008) conrmed that super hybrid rice exhibited a greater ability to produce more dry matter and its high yield
mainly resulted from the increase in dry matter accumulation after
the elongation period. The higher leaf area index and longer leaf
area duration contributed to high dry matter production and accumulation. It is necessary to conrm whether new IRRI-bred GSR
cultivars also possess a similar ability in dry matter production and
accumulation in their high yields.
This study was conducted to understand the difference of newly
bred GSR cultivars concerning biomass accumulation, allocation,
and yield formation in continuously ooded and drought-stress
conditions in comparison with drought-tolerant and irrigated rice
cultivars developed from conventional breeding systems.

2. Materials and methods

2.1. Field experiment
This study was carried out during the dry seasons of 2011

and 2012 in the experimental farms of IRRI, Los Banos,

Laguna,
Philippines. Five GSR cultivars with varying levels of tolerance
of drought were selected and compared with drought (DC) and
irrigated check (IC) varieties (Table 1). These cultivars were primarily chosen in order to understand how selection and use of the
given breeding strategy can inuence drought tolerance. They were
found to show favorable response to drought in earlier eld trials.
Details of the breeding approach adopted to develop the cultivars
are shown in Table 1. The experimental cultivars were subjected
to continuously ooded (CF) and water-limited conditions under
puddled and bunded soils. The former was in a lowland farm with
5 cm standing water until 2 weeks before harvest, and the latter
was in an upland farm where drought was applied by withdrawal
of irrigation water immediately after 30 days of transplanting, or
approximately before all test cultivars reached panicle initiation
(PI), until harvest.
A randomized complete block design was used for the experimental plot layout, with three replicates per cultivar for the 2011
experiment and four replicates for 2012. Each replicate plot has
an area of 30 m2 (5 m 6 m). An area of 5 m2 within each plot was
allotted for destructive sampling on the crop biomass components
at different phenological stages.

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M. Marcaida III et al. / Field Crops Research 162 (2014) 3038

Table 1
Characteristic features of GSR cultivars and checks used for CF and drought conditions.
Cultivar

Cross information

Breeding approach and

generation

Maturity (days after

sowing) under CF

Description

GSR IR1-12-D10-S1-D1

Huang-Hua-Zhan/
Teqing//Huang-Hua-Zhan

Backcross introgression
breeding BC1 F9

110

GSR IR1-5-S10-D1-D1

Huang-Hua-Zhan/
OM1723//Huang-Hua-Zhan

Backcross introgression
breeding BC1 F9

110

GSR IR1-8-S12-Y2-D1

Huang-Hua-Zhan/
Phalguna//Huang-Hua-Zhan

Backcross introgression
breeding BC1 F9

110

IR83142-B-19-B

IR06G103/IR06G113

105

Feng-Fu-Zhan (FFZ)

Feng-Si-Zhan/Fu-Qing-Zhan 4

Backcross introgression
breeding and designed QTL
pyramiding (DQP)
F2 pedigree breeding

Aromatic, high-yielding,
irrigated cultivar with
tolerance of salinity and
drought
High-yielding, irrigated
cultivar with tolerance of
salinity and drought
High-yielding, irrigated
cultivar with tolerance of
salinity and drought
Drought-tolerant pyramiding
line

115

Irrigated check: PSBRc82

Drought check: IR74371-70-1-1

IR47761-27-1-3-6/IRRI 108
IR55419-4/IR73278

F2 pedigree breeding
F2 pedigree breeding

110
105

In the experiment, 26- and 29-day-old seedlings were transplanted to a puddled main eld under CF (lowland) and drought
(upland) treatments in 2011, respectively. For 2012, the seedlings
were 26 and 24 days old, for CF and drought plots, respectively.
The row and plant spacing was 20 cm 20 cm with a single plant
per hill. A separate trap replicate was made in the 2011 drought
experiment, in which the third replicate of the test cultivars was
sown a week later to ensure that the effect of drought during
owering is captured fully and evenly for all varieties with oneweek growth duration differences and not allowing drought-stress
avoidance to occur. Early owering genotypes by one week that
showed drought-stress avoidance were anticipated based on previous drought trials in the same site using the same checks.
Nitrogen was applied to CF plots at 160 kg ha1 with ve
splits30 kg ha1 basal; 30 kg ha1 at 18 days after transplanting
(DAT) to promote tillering; another 30 kg ha1 at 33 and 43 DAT;
and nally, 40 kg ha1 at 64 DAT, which is around PI, to promote
spikelet differentiation. The 30 kg ha1 phosphorus and 30 kg ha1
potassium fertilizer were applied with basal nitrogen. For the
drought plots, 303030 NPK fertilizer was applied for basal and
30 kg ha1 of nitrogen was applied at 18 DAT and 33 DAT to fully
match the growth needs on nitrogen.
2.2. Data collection
Five destructive measurements were undertaken at different
stages of the rice growth season: (1) 2 weeks after transplanting
(14 DAT); (2) 21 DAT; (3) PI; (4) owering, FL; and (5) physiological maturity, PM. There were 12 hills collected per replicate at each
sampling time to measure the leaf area index (LAI); biomass weight
of green leaves, dead leaves, stem and panicles; and total aboveground biomass weight. The nal grain yield was obtained during
harvest.
Tensiometers were installed 15 cm below the soil surface to
monitor soil water tension. However, because of the severity of
water stress and because the tensiometers can read only up to
70 kPa, soil water tension was estimated using the ORYZA rice
model (Fig. 1). The 2011 experiment was marked by high temperature and low average rainfall. As such, the 2011 dry season is
referred to as the severe drought treatment, at which soil water
tension ranged from 300 to 500 kPa at the reproductive stage. The
2012 dry season was the moderate drought treatment, at which
soil water tension was 100300 kPa. The average rainfall (Fig. 1) in
the upland farm during the application of drought until harvest was

CAAS-bred GSR variety for

irrigated conditions
Irrigated
Drought-tolerant variety
recently released in the
Philippines and India

1.19 and 2.15 mm day1 , with a total rainfall of 71.6 and 155.4 mm
during severe drought (2011) and drought (2012), respectively.
2.3. Data analysis
Using analysis of repeated measures in SAS, the differences in
dry weight of green leaves, dead leaves, stem, panicles, and total
above-ground biomass were veried at 5% level of signicance
among the tested cultivars. The performance of each GSR cultivar
in specic environment conditions was compared with the check
through pair-wise comparisons to verify varietal performance differences. Dry matter components were converted to percent to
verify biomass allocation trends. Analysis of repeated measures and
pair-wise comparison with the check was also employed as in the
dry weight measurements.
3. Results
3.1. Phenology
Rice plants in drought-stressed conditions during the reproductive stage have shorter growth duration from FL to PM than those
in CF conditions (Table 2). The decrease in duration of the reproductive stage positively related to drought severity, that is, a more
severe drought shortened the time from FL to PM even more.
The drought check cultivar IR74371-70-1-1 took a longer time
from FL to PM, but gave higher panicle yield during drought stress
(Table 2). This is also true for two other GSR cultivars with higher
yield in water-limited environmentsGSR IR1-5-S10-D1-D1 and
IR83142-B-19-B.
There was no drought stress during the vegetative phase
because drought was applied one week before PI. With almost the
same growth conditions in the vegetative stage, the difference in
duration from transplanting to PI was mainly because of transplanting shock resulting from the age of seedlings.
3.2. Leaf area index (LAI)
Water stress signicantly decreased LAI relative to cultivar and
drought severity during PI and FL (Table 3). With a pair-wise
comparison between each GSR cultivar and check, a signicant difference in the drought experiment was especially observed at FL,
but not during PI and PM (Table 3). Four out of ve GSR cultivars,
which exclude IR1-5-S10-D1-D1, had a signicantly lower LAI than

M. Marcaida III et al. / Field Crops Research 162 (2014) 3038

33

Fig. 1. Amount of precipitation (mm) against soil water tension (kPa) shown in the drought experiment from the start of irrigation withdrawal until harvest whereas the
shaded areas shown in days after sowing (DAS) indicate the period of owering (FL) of the test cultivars.

the check in severe drought conditions. The opposite was observed

in drought conditions, in which all the other GSRs have signicantly
higher LAI than the check, except for GSR IR1-5-S10-D1-D1.
No signicant difference in LAI was observed at PI among test
cultivars in severe drought conditions; whereas, IR83142-B-19-B
had a signicantly high LAI in drought conditions among all the
other cultivars, even signicantly higher than the drought check.
Table 2
Phenological stage intervals (in days) for the ve GSR cultivars under varying levels
of water stress.
Phenology interval (no. of days)a

Treatment

Cultivar

S-TP

TP-PI

PI-FL

FL-PM

Severe
drought
(2011)

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check: IR74371-70-1-1
Irrigated check: PSB Rc82

29
29
29
29
29
29
29

46
46
46
39
46
32
47

32
23
26
27
34
29
31

13
22
20
21
12
26
13

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check: IR74371-70-1-1
Irrigated check: PSB Rc82

24
24
24
24
24
24
24

45
40
48
49
42
48
47

29
31
27
19
34
20
31

23
21
25
24
16
24
13

CF
(2011)

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check: IR74371-70-1-1
Irrigated check: PSB Rc82

26
26
26
26
26
26
26

45
39
45
34
43
31
44

24
29
27
29
25
27
25

28
33
26
33
36
31
29

CF
(2012)

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check: IR74371-70-1-1
Irrigated check: PSB Rc82

22
22
22
22
22
22
22

44
42
40
37
43
43
35

25
25
24
23
25
25
23

26
28
31
29
28
32
31

Drought
(2012)

S-TP = sowing to transplanting; TP-PI = transplanting to panicle initiation; PIFL = panicle initiation to owering; FL-PM = owering to physiological maturity.

3.3. Partitioning of assimilates

Accumulation of biomass in CF and water-limited experiments
was not signicantly different among the test cultivars. Differences
in the partitioning of assimilates during the reproductive phase was
notable because water stress was applied thereafter, a week before
PI. As was expected, the samples in CF environments produced
greater biomass yield, having received the necessary water requirements. In 2011, total biomass from PI to PM was 4270% lower in
severe drought than that in CF plots; whereas, it was 1020% lower
in 2012 with drought stress (Tables 4 and 5).
In severe drought conditions in 2011, no signicant difference
was found in the assimilate partitioning in different plant organs
from PI to FL among the GSR cultivars (Tables 4 and 5). Pairwise comparison with check did not show a signicant difference
in biomass allocation at PI. At FL, most of the GSR cultivars had
Table 3
Leaf area index (LAI) of GSR cultivars compared with check under drought
conditions.
Treatment

Cultivar

PI

FL

PM

Severe
drought

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

1.27
1.44
1.65
1.34
1.73
0.89

1.08bc*
2.38a
1.02c*
1.48abc*
1.21abc*
2.19ab

0.26
0.31
0.52
0.11
0.39
0.18

Drought

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

1.65ab
0.65b
1.13ab
2.17a*
1.34ab
1.42ab

2.74ab*
1.93ab
2.24ab*
2.26ab*
3.07a*
2.52b

0.93*
0.46
0.32
0.24
0.90*
0.54

CF (2011
and 2012
average)

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Irrigated check

3.42
3.72
3.45
3.63
3.77
3.57

5.28
4.42
5.03
4.89
5.49
5.16

1.62ab*
1.35ab
2.13a*
0.62b
1.17ab
0.88ab

In a column, means followed by the letters a, b, and c denote signicant difference

at 5% level by LSD; whereas, means followed by the symbol * denote signicant
difference from the check at 95% probability.

34

M. Marcaida III et al. / Field Crops Research 162 (2014) 3038

Table 4
Biomass allocation (%) of GSR cultivars and checks in drought-stressed environments during the reproductive stage.
Biomass

Cultivar

2011 severe drought stress (%)

PI

2012 drought stress (%)

FL

PM

FL

PM

Panicles

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

0
0
0
0
0
0

6
12
8
14
12
14

25c*
36ab
5c*
41a
19bc*
37a

PI
0
0
0
0
0
0

17
12
11
19
18
17

45c*
52ab
27c*
52ab
43bc*
54a

Green
leaves

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

40
52
40
56
43
44

21
22
19*
17*
17*
23

11ab
8ab
16a*
4ab
12ab*
5b

35
33*
32*
33*
35
35

20
20
16
17
20
18

8a*
3ab
6ab*
1ab
6ab*
2b

Stem

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

60
48
60
44
57
56

53
43
69
42
56
43

66b
62ab
67a*
64ab
64ab
60ab

47
49
54*
51*
49
49

51b
56ab
64a*
54ab
51b
55ab

41b
37b
54a*
40b
42b
36b

Dead
leaves

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

0
0
0
0
0
0

6*
4
6*
4
6
3

12b*
13ab
11ab*
13a
12ab
15ab

18
18*
15
17
16
16

11
12
8
11
10
10

7
7
12*
7
9
8

In a column, means followed by the letters a, b, and c denote signicant difference at 5% level by LSD; whereas, means followed by the symbol * denote signicant difference
from the drought check at 95% probability.

a signicantly higher stem biomass, except for GSR IR1-12-D10S1-D1, which had no signicant difference with the check. Dead
leaf biomass of all the GSR cultivars was signicantly higher than
that of the drought check. Trap replicates experienced the severe
drought before reaching FL that resulted to a longer duration for PI
to FL and lower biomass. Some varieties in this replicate even failed
to reach grain lling.
In the 2012 drought experiment, there was a signicant difference between the amount and percentage of assimilates allocated

to the stem and green leaves, but no signicant difference on panicle biomass at FL (Table 4). Signicant differences were noticeable
during FL, in which leaf and stem assimilates of the GSR cultivars were almost always higher than IR74371-70-1-1, the drought
check. Only GSR IR1-5-S10-D1-D1 showed no signicant difference
in stem and dead leaf biomass as compared with the check. For GSR
IR1-8-S12-Y2-D1 and FFZ, green leaf and stem biomass were higher
than the check and comparable to other GSR cultivars. On the other
hand, panicle biomass was lower at PM, indicating failure to deliver

Table 5
Dry matter accumulation and partitioning (kg ha1 ) of GSR cultivars at the reproductive stage under drought, severe drought, and CF conditions.
Cultivar/treatment

PI (kg ha1 )
GL

FL (kg ha1 )
DL

ST

PAN

TB

GL

PM (kg ha1 )
DL

ST

PAN

TB

GL

DL

ST

PAN

TB

209*
172*
229*
204*
285*
88

2208
2852*
2466*
2987*
2805*
1946

207
565
281
671
531
468

3343
4586
3658
4647
4388
3269

350ab
352ab
748a*
178b
520ab*
179b

388
592
504
612
518
559

1702b
1928ab
3263a*
1954ab
2409ab
1672b

795ab*
1606a
248b*
1916a
829ab*
1437a

3235
4478
4763
4660
4276
3846

645*
568
677*
615*
686*
391

3013bc*
2740bc
5196a*
3156bc*
3399b*
2148c

997
581
905
1086
1221
685

5855abc*
4874bc
8104c*
5837abc*
6626ab*
3938c

524a*
221ab
469ab*
73b
396ab*
133ab

465b
477b
880ab*
446b
543ab
432b

2734b
2376b
3945a*
2555b
2627b
2048b

3026ab
3350a
1956b*
3328a
2686ab
3012ab

6748
6425
7250*
6401
6252
5625

Severe drought
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

653
641
661
590
922*
504

0
0
6
0
0
0

975
582
996
473
1230
638

0
0
0
0
0
0

1628
1223
1663
1063
2151
1141

Drought
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

902
645
725
1069*
699
731

451
362
334
538*
321
339

1215
975
1222
1648*
996
1034

0
0
0
0
0
0

2568
1982
2282
3255*
2016
2103

CF (2011)
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Irrigated check

1349
1377
1667
812
1313
1421

6
7
12
8
9
15

1571
1606
2040
1080
1495
1499

0
0
0
0
0
0

2926
2990
3719
1899
2817
2935

2862
3165
3073
2404*
3082
3003

38
35
81
51
50
63

4541
4876
5165
4431*
5580
4617

1415
1723
1326
1483
1784
1650

8855
9800
9644
8369
10496
9333

1859ab*
1548ab
1925a*
856b
1696ab*
1112ab

660b*
872ab*
653b*
964ab*
930ab*
1470a

4622
4105
5012*
3963
4893*
4026

7249a
7019a
5040b*
7050a
7925a*
6741a

14,389
13,543
12,630
12,833
15,445*
13,349

CF (2012)
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Irrigated check

920
1112
1292
1093
898
1114

0
0
0
0
0
0

1229
1508
2184
1967
961
1352

0
0
0
0
0
0

2149
2620
3476
3060
1859
2466

1689
1334
1891
1550
1714
1694

769*
808
961
755*
889
903

3323
3193
4463
3336
3208
3381

1119
1345
1602
843*
1217
1322

6900
6680
8917
6483*
7028
7299

91
134*
76
54
36*
70

1477*
1591*
2247*
1503*
1723*
858

1192
1201
1500*
936
1473
949

5418a
5106a
5770b*
4852a
5880a*
4599a

8178
8032
9593*
7345
9111*
6476

719
997
682
785
768
767
1200a*
985ab*
1326a*
980ab*
1321a*
713b

Assimilate partitions: GL = green leaves, DL = dead leaves, ST = stem, PAN = panicles, and TB = total biomass. In a column, means followed by the letters a, b, and c denote
signicant difference at 5% level by LSD; whereas, means followed by the symbol * denote signicant difference from drought check at 95% probability.

M. Marcaida III et al. / Field Crops Research 162 (2014) 3038

carbohydrates to the storage organ or a lower transferring capability of reserved soluble carbohydrates from stem to panicle from FL
to PM.
There was no signicant difference in the total biomass during
PM across cultivars in two drought environments. Under CF conditions, GSR IR1-8-S12-Y2-D1 and FFZ gave a signicantly higher
total biomass than all other cultivars.
Upon reaching PM, IR83142-B-19-B and GSR IR1-5-S10-D1-D1
showed more promising performance than other GSR cultivars,
with consistently high panicle yield in both CF and water-stressed
environments (Tables 4 and 5). Both cultivars gave a panicle yield
of at least 1.5 tons ha1 and 35% dry mass allocated to panicles in
severe drought conditions (2011), which was 1233% higher than
the drought check. In the 2012 drought conditions, the panicle yield
of both cultivars was at least 3.3 tons with dry mass allocation to
the panicles 52% of the total above-ground biomass (TB). Panicle
biomass thaw was 1011% higher than that of the drought check.
Panicle yield under CF conditions ranged from 5 to 7 tons ha1 , with
panicle biomass of 5266% of TB. By ANOVA and pair-wise comparison, however, the advantage in panicle biomass shown by these two
GSR cultivars was not signicant compared to the check varieties
(Table 5).
Plant materials with low percentage of green leaf biomass at PM
have relatively higher proportions of panicle biomass as in the case
of IR83142-B-19-B and GSR IR1-5-S10-D1-D1 (Table 4). The opposite is true for GSR IR1-8-S12-Y2-D1, which has the highest total
above-ground biomass at PM in the two drought environments,
but has the lowest panicle yield among all the other GSR cultivars. These GSR cultivars, including GSR IR1-12-D10-S1-D1 and FFZ,
have a high amount and percentage of assimilates in green leaves
and stem, indicating their failure to efciently deliver the reserved
soluble carbohydrates to the reproductive organs in water-limited
conditions (Tables 4 and 5). The shorter duration from FL to PM,
implying short duration for grain lling, also contributed to lowyielding GSR cultivars under drought stress (Tables 2 and 5).
Grain yield was measured upon harvest and shown in Table 7.
Yield in irrigated plots ranged from 5 to 7.9 tons ha1 in 2011
and from 6.6 to 11.5 tons ha1 in 2012. As observed at PM in
both drought environments, cultivars in severe drought stress gave
lower grain yield (0.281.67 tons ha1 ) than those in moderate
drought stress (1.282.91 tons ha1 ), with IR83142-B-19-B and GSR
IR1-5-S10-D1-D1 consistently showing higher yield than the other
cultivars. In comparison with the check varieties, grain yield of
GSR cultivars was higher in the 2011 irrigated environment. The
opposite was observed in 2012, in which the check variety had
higher grain yield than all the GSR cultivars except for GSR IR1D10-S1-D1. In the water-limited environment, results showed that
the check variety had higher grain yield than all the GSR varieties
under severe drought conditions (2011). While GSR IR1-5-S10-D1D1 and IR83142-B-19-B have a relatively higher grain yield than
the check, all the other GSR cultivars have close but lower grain
yield than the check.

4. Discussion
In the dry seasons of 2011 and 2012, drought environments
were created after irrigation was stopped. Slow elongation of
stems, steep leaf angle, short plant height, and leaf-rolling, which
manifested in the decreased LAI and biomass weight, were
observed more in the GSR cultivars under drought than under CF
(Tables 3 and 5). These indicate that the GSR cultivars have similar drought coping mechanisms as commonly observed for plants
subjected to drought stress to avoid dehydration and complete
their life cycle (Wopereis et al., 1996; Chaves et al., 2003). The crucial and severe effect of drought stress in rice happens during the

35

reproductive and grain-lling stages while reserved carbohydrates

in vegetative organs are translocated to the panicles. Boonjung and
Fukai (1996) reported that the inability of leaves and stems to effectively deliver stored carbohydrates to the storage organs causes low
yield.
Since no signicant difference was observed in all the experimental plots during the vegetative stage when drought stress was
not yet applied, the discussions herein focus mainly on the partitioning of assimilates from PI to PM. The differences in assimilate
partitioning and growth-stage duration dened the ability of GSR
to cope with drought stress as shown by the panicle yield of the
cultivars.
There was an obvious decline in biomass yield compared to CF
treatments in the reproductive stage when drought was applied. It
is necessary to examine the biomass accumulation and assimilate
partitioning behavior that enabled the well-performing GSR cultivars to cope with stress brought about by limited water availability
and still produce high panicle and grain yield (Tables 4 and 7).
To examine the behavior of the GSR cultivars together with the
drought check, biomass production rate analysis was employed.
The rate of biomass production (kg ha1 day1 ) is an estimate
of the amount of biomass productivity between two phenological
stages. It is computed by getting the difference in biomass weight
between two phenological stages divided by the stage duration (in
days). A positive value represents an increase in the rate of biomass
formation, whereas a negative value indicates a decrease in the rate
of biomass production resulting from leaf death or translocation of
carbohydrates to the panicles and the burning of carbohydrates
to survive the drought (Table 6). As in the study of Boonjung and
Fukai (1996), variation in biomass produced during specic growth
stages affects the grain yield.
When the rice plants reach FL, the carbohydrates stored in the
vegetative organs are delivered to the panicles until PM. Therefore, it is important to realize that the biomass produced in the
leaves and stem from the vegetative stage to PI is needed to build
up enough sources for translocation. Table 6 shows the translocation (by the vegetative organs) and accumulation (by the storage
organs) of assimilates from FL to PM. It is noticeable how biomass
in the leaves and stem drastically decreased from FL to PM in the
drought experiments. This mechanism is indicative of the plants
coping strategy at the sense of water stress, thereby producing
reserves to be able to mobilize itself for panicle production during
grain lling (Barnabas et al., 2008; Chaves et al., 2003; Yang and
Zhang, 2006). Efcient translocation is indicated by the high accumulation rate of the panicle and more negative values for the leaves
and stem (Table 6). The negative production rates of the stem and
leaves of GSR IR1-12-D10-S1-D1, GSR IR1-5-S10-D1-D1, IR83142B-19-B, and FFZ indicate delivery of carbohydrates to the panicles
from FL to PM. The opposite was observed for GSR IR1-8-S12-Y2D1, in which it continued to accumulate reserves in the vegetative
organs but failed to translocate them, thus it has the lowest panicle and grain yield in severe drought stress (2011). Its panicle and
grain yield were also the lowest in moderate drought stress (2012)
because of low translocation coefcient, although reserves were
used for grain lling (Tables 5 and 7).
Comparing the delivery of carbohydrates during PI to FL
(Table 6) showed a rate of assimilate accumulation almost similar in the leaves and stem during PI to FL in the 2011 and 2012
experiments. The drought experiment in 2012 showed that GSR
cultivars can produce a panicle yield of at least 2 tons ha1 (Table 5)
under moderate drought (100300 kPa) through their efciency in
translocating carbohydrates to the panicles from FL to PM.
As shown in studies by Yang and Zhang (2006) and Boonjung and
Fukai (1996), the time to reach maturity was shortened as observed
in poorly performing GSR cultivars (Table 2), and it was very apparent in severe drought. Water stress during grain lling could hasten

36

M. Marcaida III et al. / Field Crops Research 162 (2014) 3038

Table 6
Rate of biomass accumulation in ve GSR cultivars under three different crop growth conditions.
Cultivar/treatment

PI-FL (kg ha1 day1 )

FL-PM (kg ha1 day1 )

TL

ST

PAN

TB

TL

ST

PAN

TB

Severe drought
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

8
24
10
15
3
12

38
100
57
93
46
45

6
25
11
25
16
16

52
149
78
133
63
73

15
10
17
10
1
5

40
41
41
49
34
11

46
47
2
59
26
37

9
4
56
0
9
21

Drought
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Drought check

17
18
36
1
29
2

62
58
150
79
71
56

34
19
34
57
36
34

113
95
220
135
136
92

38
40
27
45
69
22

12
17
51
25
50
4

89
130
42
93
94
97

39
73
36
23
25
71

CF (2011)
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Irrigated check

63
63
55
55
73
66

122
113
116
114
163
126

58
59
49
51
71
67

243
235
220
220
307
259

14
24
22
19
14
16

3
24
6
14
19
20

208
162
143
170
171
174

197
114
115
137
138
138

CF (2012)
GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Irrigated check

60
42
65
53
69
66

83
68
95
59
90
90

44
54
67
36
49
59

187
164
227
148
208
215

34
14
18
26
31
53

82
71
96
83
63
78

165
133
134
139
170
106

45
48
20
30
76
25

Assimilate partitions: TL = leaves (green + dead), ST = stem, PAN = panicles, TB = total biomass. Phenological stage intervals: PI-FL = panicle initiation to owering, FLPM = owering to physiological maturity.

maturity. In favorable and non-resource-limited environments, the

time between owering to maturity takes at least 30 days; however, it was cut short into at least 12 days in severe drought and
16 days in drought conditions. On the contrary, the drought check
together with GSR IR1-5-S10-D1-D1 and IR83142-B-19-B took a
longer time (2126 days), allowing them to have a long period for
translocation of carbohydrates and for generation of assimilates to
facilitate grain lling. In contrast, GSR IR1-12-D10-S1-D1 and FFZ
tried to cope with drought stress by speeding up maturity, but were
unsuccessful in delivering as much assimilates to the panicles. This
lowered translocation efciency relative to those under CF, thus the
low panicle yield at PM.
Among the ve GSR cultivars used in this study, GSR IR1-5S10-D1-D1 and IR83142-B-19-B were able to withstand different
levels of water stress with 1.63.3 tons ha1 of panicle yield at PM
(Table 5), and 1.32.6 tons ha1 of grain yield at harvest. These two
cultivars distinctly had the ability to effectively translocate assimilates stored in the vegetative organs from previous growth stages
(Table 6). Furthermore, high translocation was facilitated by not
having to dramatically cut short their grain-lling duration, unlike
with other cultivars (Table 2).

Under severe drought, IR83142-B-19-B was able to sustain its

photosynthetic activity and effectively translocate carbohydrates
to the panicles as manifested by the lack of signicant change in
TB from FL to PM (Table 5). Low LAI under moderate drought stress
(Table 3) allowed low water requirements for transpiration and
photosynthesis; hence, photosynthesis is more effective per unit
area of LAI with no signicant difference in TB among GSR cultivars. In the case of GSR IR1-5-S10-D1-D1 under severe drought, in
which LAI is relatively higher (Tables 3 and 5), the product of photosynthesis was adequate for plant maintenance. Thus, there was
no signicant change in TB from FL to PM (i.e., growth rate of TB
almost 0). Both cultivars (IR83142-B-19-B and GSR IR1-5-S10-D1D1), with enough carbohydrates produced from photosynthesis,
have a very efcient translocation ability that enabled them to
cope with different drought-stress conditions, and this is why they
performed better than the other GSR cultivars.
Comparing panicle yield at PM with the drought check under
moderate drought stress, there was no signicant difference
observed in four GSR cultivarsGSR IR1-12-D10-S1-D1, GSR IR15-S10-D1-D1, GSR IR1-8-S12-Y2-D1, and FFZ (Table 5). These
cultivars are able to produce a panicle yield of 3.3 tons ha1

Table 7
Grain yield of GSR cultivars and checks at harvest.
Cultivar

GSR IR1-D10-S1-D1
GSR IR1-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Irrigated check
Drought check

Irrigated

Drought

2011

2012

2011

2012

7.25 (+7.6)
7.02 (+4.2)
5.04 (25.2)
7.05 (+4.6)
7.93 (+17.7)
6.74
5.78

11.48 (+1.5)
10.35 (8.5)
10.53 (6.9)
10.52 (+7.0)
10.74 (5.0)
11.31
6.56

0.46 (74.3)
1.30 (27.4)
0.28 (84.3)
1.67 (6.7)
0.81 (54.7)
1.29
1.79

2.03 (2.4)
2.60 (+25.0)
1.28 (38.5)
2.91 (+40.0)
1.98 (4.8)
2.68
2.08

The gures in parenthesis are yield differences relative to the check (i.e., irrigated check under irrigated conditions and drought check under drought-stressed conditions).

M. Marcaida III et al. / Field Crops Research 162 (2014) 3038

37

Table 8
Multi-environment and multi-treatment yield of GSR cultivars in DS 2011 and 2012.
Cultivars

GSR IR1-12-D10-S1-D1
GSR IR1-5-S10-D1-D1
GSR IR1-8-S12-Y2-D1
IR83142-B-19-B
FFZ
Irrigated check
Drought check
a
b
c

Seasonal meana (tons ha1 )

Mean by treatmentb (tons ha1 )

2011

2012

Irrigated

Drought

3.86
4.16
2.66
4.36
4.37
4.02
3.79

6.76
6.48
5.91
6.72
6.36
7
4.32

9.37
8.69
7.79
8.79
9.34
9.03
6.17

1.25
1.95
0.78
2.29
1.4
1.99
1.94

Varietal mean across seasons

and treatmentsc (tons ha1 )

Percent advantage over

drought check

5.31
5.32
4.28
5.54
5.37
5.51
4.05

31.1
31.4
5.7
36.8
32.6
36.0

Seasonal mean is the average grain yield of the irrigated and drought treatments combined for each season.
Mean by treatment is the average grain yield for each treatment during the two experimental seasons.
Varietal mean is the average grain yield of each cultivar with treatments and seasons combined.

and grain yield of 2.62.9 tons ha1 at a soil water tension of

100300 kPa, hence, they can be recommended for drought-prone
environments.
When compared to the drought check, the grain yield of
IR83142-B-19-B and GSR IR1-5-S10-D1-D1 was lower in severe
drought and not signicantly higher in moderate drought environments. However, these two cultivars continue to show yield
advantage over the other GSR samples.
Resilience of GSR cultivars could be ascertained by making an
effective comparison on (1) the mean grain yield across seasons
(DS 2011 and 2012), from which the average of the irrigated and
drought experiments were obtained; (2) the treatment (irrigated
and drought), in which the 2-year means of each treatment was
computed; and (3) the mean varietal performance across seasons
and treatments (Table 8). Regardless of the variability of seasonal
weather and the availability of water, the cultivars GSR IR1-5S10-D1-D1 and IR83142-B-19-B were fairly consistent in obtaining
high grain yield of 2.08.8 tons ha1 and were comparable and/or
higher than the checks. In typical rainfed lowland situations in Asia,
drought does not recur each year with the same intensity; some
years may have favorable seasons. Therefore, varieties need to be
more resilient and should have a relatively high yield in all such
conditions (Latte et al., 2006).
For the yield average of the drought and irrigated experiments
in 2011 (Table 8), when the drought stress was more severe, all the
GSR varieties gave a yield of at least 2.5 tons ha1 . Three cultivars
gave higher yield than the irrigated and drought checks. In 2012,
all the GSR varieties have a higher yield than the drought check,
giving a combined average yield from irrigated and drought plots
of 5.96.7 tons ha1 .
The mean of each treatment from two seasons (Table 8) only
afrm earlier ndings about the varieties that perform well and
the ability of GSR to produce high yield in non-water-limited environments.
The multi-environment mean in Table 8 showed the resiliency
of GSR cultivars, having been exposed to two dry seasons in two different environments (water-limited and irrigated). While they gave
almost the same yield as the irrigated check, all the grain yields of
the GSR cultivars, which ranged from 4.3 to 5.5 tons ha1 , were at
least 5.736.7% higher than the drought check. This indicates that,
given the unpredictability of weather conditions in rice-producing
areas every year, in the presence or absence of drought, GSR cultivars are capable of producing high grain yield in the long run.
5. Conclusions
With the introduction of water stress before the onset
of the reproductive stage, GSR entries IR1-5-S10-D1-D1 and
IR83142-B-19-B were able to withstand drought of varying levels, thereby producing grain yield of 1.12.9 tons ha1 . These two

drought-tolerant GSR cultivars did not have to abruptly shorten

their duration from FL to PM, compared with other GSR test materials, to reduce the grain-lling period in order to cope with water
limitation. This could have allowed efcient translocation of assimilates and reserved carbohydrates from the vegetative organs to the
ultimate sink (panicle yield). Low LAI enabled balanced biomass
accumulation with low transpiration, preventing signicant reduction in grain lling.
GSR tolerant cultivars subjected to 100300 kPa of water stress
were able to deliver more products of photosynthesis and stored
carbohydrates to the panicles than those under severe water stress,
and were able to produce 2.03.3 tons ha1 of panicle yield.
This study also showed that introgression breeding employed
in GSR was successful in developing drought-tolerant cultivars.
Although not expressed well by all the GSR cultivars under severe
drought conditions (soil tension higher than 300 kPa at 15-cm soil
depth), those cultivars that did not perform well under severe
drought stress could have salinity and ooding tolerance besides
moderate drought tolerance and there exists a compensation
mechanism when materials are selected for multiple abiotic stress
conditions. Grain yield of at least 2 tons ha1 could be expected
from GSR cultivars in moderate drought conditions with soil water
tension of 100300 kPa at a 15-cm depth. Multi-environment comparison also showed the resiliency of the GSR cultivars, thereby
producing an average grain yield of 4.35.5 tons ha1 in two dry
seasons, with yields from drought and irrigated environments combined.
Acknowledgments
The funds for this research were provided under the Green
Super Rice Project GD1393-1 by Bill & Melinda Gates Foundation
through Chinese Academy of Agricultural Sciences (CAAS), Beijing
to the International Rice Research Institute, Philippines.
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