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Agriculture, Ecosystems and Environment 122 (2007) 233–242

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Impact of elevated CO2 and temperature on rice yield and methods
of adaptation as evaluated by crop simulation studies
P. Krishnan a,*, D.K. Swain b, B. Chandra Bhaskar b, S.K. Nayak a, R.N. Dash b
a

Division of Biochemistry, Plant Physiology and Environmental Sciences, Central Rice Research Institute, Cuttack 753006, India
b
Division of Soil Science and Microbiology, Central Rice Research Institute, Cuttack 753006, India
Received 7 February 2006; received in revised form 20 December 2006; accepted 19 January 2007
Available online 6 March 2007

Abstract
Impact of elevated CO2 and temperature on rice yield in eastern India was simulated by using the ORYZA1 and the INFOCROP rice
models. The crop and weather data from 10 different sites, viz., Bhubaneswar, Chinsurah, Cuttack, Faizabad, Jabalpur, Jorhat, Kalyani, Pusa,
Raipur and Ranchi, which differed significantly in their geographical and climatological factors, were used in these two models. For every
1 8C increase in temperature, ORYZA1 and INFOCROP rice models predicted average yield changes of 7.20 and 6.66%, respectively, at
the current level of CO2 (380 ppm). But increases in the CO2 concentration up to 700 ppm led to the average yield increases of about 30.73%
by ORYZA1 and 56.37% by INFOCROP rice. When temperature was increased by about +4 8C above the ambient level, the differences in the
responses by the two models became remarkably small. For the GDFL, GISS and UKMO scenarios, ORYZA1 predicted the yield changes of 
7.63, 9.38 and 15.86%, respectively, while INFOCROP predicted changes of 9.02, 11.30 and 21.35%. There were considerable
differences in the yield predictions for individual sites, with declining trend for Cuttack and Bhubaneswar but an increasing trend for Jorhat.
These differences in yield predictions were mainly attributed to the sterility of rice spikelets at higher temperatures. Results suggest that the
limitations on rice yield imposed by high CO2 and temperature can be mitigated, at least in part, by altering the sowing time and the selection
of genotypes that possess higher fertility of spikelets at high temperatures.
# 2007 Elsevier B.V. All rights reserved.
Keywords: Climate change; CO2; INFOCROP; ORYZA; Oryza sativa L.; Simulation; Temperature; Yield

1. Introduction
The climatic variability and the predicted climatic changes
are of major concern to the rice crop scientists because of
Abbreviations: BVP, Basic Vegetative Phase; DLAI, Death Rate of
Leaf Area Index; FACE, Free Air Concentration Enrichment; GCMs,
General Circulation Models; GFDL, General Fluid Dynamics Laboratory;
GFP, Grain Filling Phase; GISS, Goddard Institute of Space Studies; GLAI,
Leaf Area Growth Rate; IPCC, Intergovernmental Panel on Climate
Change; LAI, Leaf Area Index; NATP, National Agricultural Technology
Project; PFP, Panicle Formation Phase; PLTR, Net loss of LAI due to
transplanting; PSP, Photoperiod-Sensitive Phase; RLAI, Net Leaf Area
Growth Rate; RUE, Radiation Use Efficiency; RWLVG, Increment in Leaf
Weight; SLA, Specific Leaf Area; SUBLAI, LAI is simulated in the
subroutine SUBLAI; UKMO, United Kingdom Meteorological Office
* Corresponding author.
E-mail address: prameelakrishnan@yahoo.com (P. Krishnan).
0167-8809/$ – see front matter # 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.agee.2007.01.019

their potential threat to rice productivity and the associated
impact on the socioeconomic structure of many rice-growing
countries. Among the global atmospheric changes, the increasing concentrations of greenhouse gases such as CO2 may
have significant effect on rice productivity due to increase in
both the average surface temperature and the amount of CO2
available for photosynthesis (Aggarwal, 2003). In the absence
of temperature increase, many studies have shown that the net
effect of doubling of CO2 was increase in the yield of rice
(Kim et al., 2003). It becomes necessary to assess the effects of
potential interactive changes of CO2 and temperature in order
to determine the future agricultural strategies that would
maintain higher rice productivity.
The simulations by different models and many field
experiments have shown the potential impact of climatic
change and the variability in rice productivity (Baker et al.,

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P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

1992b; Peng et al., 2004; Kim et al., 2003). The modeling
studies from Bangladesh (Karim et al., 1994), Japan (Horie
et al., 2000), China (Bachelet et al., 1995) and India (Mall
and Aggarwal, 2002) reported the country-wise variations in
rice production, anticipated due to the climatic changes. The
simulated yields increased when temperature increases were
small, but declined when the decadal temperature increase
was more than 0.8 8C, with the greatest decline in crop
yields occurring between the latitudes of 108 and 358N.
Similar results were obtained by Penning de Vries (1993).
Many uncertainties exist in modeling studies, partly due
to the quality of the predictions by the models, from the use
of limited sites for which historical weather data are
available, due to the quality of the crop simulation models,
especially when applied under the rain-fed conditions
(Bachelet et al., 1995), and due to the quality of the climate
models used to predict future weather scenarios. These
uncertainties may be reduced only when a large number of
scenarios for different locations are compared and evaluated.
In order to overcome the predicted limitations for rice
production in the future, there is also a need to identify and
evaluate the suitable agronomic practices such as altered
sowing date and selection of improved varieties with
increased spikelet fertility at high temperature and other
useful traits.
Attempts have been made earlier to assess the general
effects of global environmental changes on rice yield using
simulation models, but little attention has been given to the
potential interactive effects between high temperature and
increasing CO2 levels. Lal et al. (1998) used the CERES rice
and predicted a 20% decline in rice yields in the
northwestern India due to elevated CO2 and temperature.
Eastern India accounts for about 63% (26.5 million ha) of
the total rice-growing area in India. The rice ecosystems in
these regions show characteristic differences with respect to
the environmental factors as well as the cultural practices.
There is an urgent need to characterize the impact of future
climatic changes on rice yield in these regions for sustaining
the productivity. The present paper discusses the outcomes
of two rice growth simulation models, ORYZA and
INFOCROP rice, when applied to different rice-growing
sites in the eastern India for studying the potential interactive
effects between high temperature and increasing CO2 levels
and for the different thermal climate change scenarios.

2. Materials and methods
Two popular models of rice growth ORYZA1 (Kropff
et al., 1994) and INFOCROP rice (Aggarwal et al., 2006) are
used in this study. Prior to their use, both were evaluated
and compared (Fig. 1) and then these crop models were
calibrated for the indica variety IR 36 at all sites. In general,
the two models differ in the manner in which dry matter
production and partition, leaf area development, and phenological development are calculated. ORYZA1 calculates dry

Fig. 1. Comparison of yield simulated by ORYZA1 and INFOCROP rice.

matter production as a function of light, CO2 and temperature
by considering photosynthetic processes at the leaf level and
integrating these over the canopy to obtain crop-level values.
Respiration is also modeled explicitly as a function of
temperature and partitioning of dry matter is according to
phenology-dependent functions. Thus, ORYZA calculates
dry matter production as a function of gross canopy
photosynthesis, depending on the detailed calculations of
the distribution of light within the canopies, the radiation
absorbed by the canopy and photosynthesis light response
curve of leaves. Growth and maintenance respiration are
calculated as a function of tissue N content, temperature
and crop-specific coefficients. This methodology, although
yields very accurate results, poses practical difficulties
because of its requirement for detailed and careful measurements (Kropff et al., 1994). The INFOCROP model, however,
uses a simpler radiation use efficiency (RUE) relationship
between intercepted solar radiation and growth, in which
respiration is implicit. CO2 effects are accounted for by using
a curvilinear function relating RUE to CO2 concentration.
Temperature is assumed not to affect RUE. More or less
similar results can be obtained under normal radiation
situations by calculating the net dry matter production as a
function of RUE. Pre-determined values of RUE were input in
the model as a function of crop/cultivar (RUEMAX), and
RUE was further modified by the development stage
(Aggarwal et al., 2004).
The rice-growing regions included for the present study lie
in eastern India, and these sites (Bhubaneswar, Chinsurah,
Cuttack, Faizabad, Jabalpur, Jorhat, Kalyani, Pusa, Raipur
and Ranchi), which are geographically apart at varied
altitudes ranging from 7.8 to 86.56 m above mean sea level,
show characteristic features with respect to the weather and
crop factors. The crop parameters included in the study were
obtained from the field experiments conducted at these sites
under National Agricultural Technology Project (NATP)
RRPS 25 during 2001–2003 (Table 1).
The input parameters used for the two models are given in
Table 2a (parameters such as varietal data, soil data and
weather data), Table 2b (physical and chemical properties of

P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242
Table 1
Sites and institutes where the field observation and weather data were
collected
Sites

Institutes

Bhubaneswar
Chinsurah
Cuttack
Faizabad

Orissa University of Agriculture & Technology (OUAT)
Rice Research Station (RRS)
Central Rice Research Institute (CRRI)
Narendra Deva University of Agriculture &
Technology (NDUAT)
Jawaharlal Nehru Krishi Vishwa Vidyalaya (JNKVV)
Assam Agricultural University (AAU)
Bidan Chandra Krishi Viswa Vidyalaya (BCKV)
Rajendra Agricultural University (RAU)
Indira Gandhi Agricultural University (IGAU)
Birsa Agricultural University (BAU)
Indian Meteorological Department (IMD),
Pune (weather data)

Jabalpur
Jorhat
Kalyani
Pusa
Raipur
Ranchi

235

were made for the main rice-growing season for each of
these sites. Generally, the dates of sowing and transplanting
were supplied along with the weather data.
2.1. Climate change scenarios
For each of these rice-growing regions, the potential yield
of rice was simulated under 30 different combinations of
CO2 and temperature, including with the ‘fixed increment’
changes in CO2 (380, 400, 500, 600 and 700 ppm) and
temperature (ambient, +1, +2, +3, +4 and +5 8C) individually, and with all the combinations of these levels of CO2 and
temperature. The actual daily weather data, collected for
three consecutive years from each of these sites were used
for simulation. For temperature changes, the daily weather
data from each site were exported to Excel spread sheets and
then the daily maximum and minimum temperatures were
increased by 1–5 8C individually. Later, the modified
weather data were used as the inputs for ORYZA1. In case
of INFOCROP, for a given daily weather data, the climate

soils) and Table 2c (geographical information). The weather
data containing measurements of sunshine hours, temperature and rainfall from all these sites were used as the baseline
data for input into these crop models. Later, the simulations
Table 2a
Combined input parameters for both the models used
Varietal data

Soil data

Weather data (daily data)

1. Base temperature for sowing to germination

1. Soil type

2. Thermal time for sowing to germination
3. Base temperature for germination to 50% flowering
4. Thermal time for germination to 50% flowering
5. Base temperature for 50% flowering to physiological maturity
6. Thermal time for 50% flowering to physiological maturity
7. Optimal temperature
8. Maximum temperature
9. Sensitivity to photoperiod
10. Relative growth rate of leaf area
11. Specific Leaf Area
12. Extinction coefficient of leaves at flowering
13. Radiation Use Efficiency
14. Root growth rate
15. Index of greenness of leaves
16. Sensitivity of crop to flooding
17. Index of N fixation
18. Slope of storage organ number/m2 to dry matter during storage
organ formation stage
19. Potential storage organ weight at maximum temperature
20. Nitrogen content of storage organ
21. Sensitivity of storage organ setting to low temperature
22. Sensitivity of storage organ setting to high temperature

2. Soil texture
3. pH of soil
4. EC
5. Sand %
6. Silt %
7. Clay %
8. Saturation fraction
9. Field capacity
10. Wilting point
11. Saturated hydraulic conductivity
12. Bulk density
13. Organic carbon

1. Maximum and minimum
temperatures
2. Relative humidity
3. Sun shine hours
4. Precipitation

Table 2b
Model inputs of physical and chemical properties of soil at different locations
Soil properties

Bhubaneswar

Kalyani

Faizabad

Ranchi

Pusa

Raipur

Jabalpur

Jorhat

Cuttack

Texture
B.D.a (g/cc)
O.C.b (%)
Total N (%)
Available N (kg/ha)

Sandy clay loam
1.35
0.73
0.07

Sandy clay loam
1.45
0.51
0.04

Loam
1.3
0.3

Loam
1.41
0.68

Sandy clay loam
1.3
0.57

Clayey
1.34
0.69

Sandy clay loam
1.48
0.67

Sandy loam
1.4
0.47
0.07

128.4

355

258.6

Silt
1.23
0.54

267

265.8

a
b

B.D., bulk density.
O.C., organic carbon.

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P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

Table 2c
Model inputs of geographical information of different locations
Soil properties
Latitude
Longitude
Altitude (m)

Bhubaneswar
0

20814 N
858520 E
25.9

Kalyani
0

22857 N
888210 E
7.8

Faizabad
0

22887 N
88840 E
8.62

Ranchi
0

23817 N
858190 E
625

change scenarios were specified as +1 to +5 8C for
temperatures and the percent changes in precipitation.
The inbuilt weather generator of that model generated the
modified daily weather data accordingly, and these data were
used as the daily weather inputs. The climate change
scenarios under different levels of CO2 were applied by
changing the ambient CO2 concentration parameter in these
two models.
Besides, the outputs from the climate models were used
in combination with these crop models. The coarse grid from
each GCM was interpolated using a four point inversedistance-squared algorithm to a 0.58 latitude  0.58 longitude grid using a raster-based Geographical Information
Systems (GIS) software package. Scenarios were produced
by applying ratios of precipitation or differences in temperature predicted for the 2  CO2 and 1  CO2 simulation to
the baseline present climate data set for different sites
(Mathews and Wassmann, 2003). The important features of
three General Circulation Models (GCMs) used such as
the General Fluid Dynamics Laboratory (GFDL) Model,
Goddard Institute of Space Studies (GISS) model and the
United Kingdom Meteorological Office (UKMO) model are
provided in Table 3.
The GCM scenarios were produced by applying the ratios
of precipitation or differences in temperature predicted for
the 1  CO2 (380 ppm) and 2  CO2 (760 ppm) simulations
to the baseline daily weather data set. The changes between
1  CO2 and 2  CO2 conditions were representative of the
differences between the present and the future climate
scenarios following an equivalent doubling of CO2.
2.2. Simulation
For the simulation analysis, only runs that terminated
normally or by crop deaths as a result of temperature were
included. The relative yield changes under the scenarios,
referred to yields predicted for the current climate, were
used in the analysis rather than absolute yields. To provide
an estimate of the overall effect of climate change on rice
production under the three GCMs scenarios, the average

Pusa

Raipur

0

0

2285 N
868E
23

25859 N
85840 E
51.84

Jabalpur
0

23809 N
798580 E
411

Jorhat
0

2688 N
95850 E
86.56

Cuttack
208300 N
868E
23

relative increase predicted for each site was weighed by its
current production observed in the field trials of an earlier
study (Annual Report of NATP RRPS 25, 2002–2003). The
differences in the production capacities among these sites
were evaluated.

3. Results
3.1. Effect of temperature and CO2 levels at fixed
increments on yields
At all the CO2 levels tested (380, 400, 500, 600 and
700 ppm), both the models predicted the declining yields of
rice due to an increase in temperature. On the contrary, an
increase in CO2 level at any particular temperature increased
the rice yields (Table 4). At the current level of CO2
(considered at 380 ppm), ORYZA predicted a mean change
of 7.20% in yields for every 1 8C increase in temperature,
while INFOCROP predicted 6.66%. But increasing CO2
concentration (700 ppm) resulted in increases of 30.37 and
56.37% in yield by ORYZA and INFOCROP, respectively.
However, with temperature increase of +4 8C above
ambient, the differences in the yield predictions by the
two models became remarkably small (Table 4).
3.2. Effect of predicted GCM scenarios on rice yields
The predicted changes in overall production for each site
under different climate scenarios using the two crop models
are provided in Tables 5a and 5b. In general, the ORYZA
model suggested the decreases of 7.63, 9.38 and 
15.86% in yield for the GDFL, GISS and UKMO
scenarios, respectively. For the corresponding scenarios,
INFOCROP indicated larger reductions at 9.02, 11.30
and 21.35%, respectively. When each site was analysed
individually under three GISS and UKMO scenarios, almost
all the sites except one showed the declining trend in yields.
The decreases in the yield of rice, when the mean of the
corresponding values in Tables 5a and 5b were considered,

Table 3
The major features of the General Circulation Models used in this study

Source laboratory
Base 1  CO2 (ppm)
Change in global temperature (8C)
Change in global precipitation (%)

GFDL

GISS

UKMO

Geophysical Fluid Dynamics
Laboratory
380
+4.0
8

Goddard Institute for
Space Studies
380
+4.2
11

United Kingdom
Meteorological Office
380
+5.2
15

P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

237

Table 4
Mean predicted changea (%) in the potential yield under the ‘‘fixed’’ temperature and CO2 scenarios
CO2 concentration

Temperature increments (8C) b
0

+1

+2

+3

+4

+5

Average

INFOCROP
380
400
500
600
700
Average

0
5.23
25.92
40.61
56.37
25.14

2.68 
2.90
13.90
29.39
41.21
16.86 

18.04 
12.03
3.60
16.74
27.96
3.65 

28.15 
23.21 
12.32
1.04
9.85 
10.56 

36.36 
33.05 
25.08 
15.80 
7.67 
23.59 

42.49 
40.32 
35.11 
29.17 
23.15 
34.05 

20.39 
18.12 
4.85
7.14
17.43 
3.76

ORYZA
380
400
500
600
700
Average

0
2.78
18.57
26.8
30.73
16.27 

4.27
0.93
11.73
18.54
23.77
10.14 

15.32 
8.44
0.24
5.03
8.67 
1.96 

26.78 
19.74 
11.75 
6.86 
3.68 
13.76 

29.45 
23.67 
15.43 
10.87 
7.32 
17.35 

38.96 
33.65 
26.57 
21.84 
19.04 
28.01 

19.13 
13.22 
3.87
1.80
5.52 
5.78

a
b

Changes are averaged across all sites and at all the available years.
Temperature increments are above the current temperatures at each site.

Table 5a
Estimated changes in rice yield predicteda by the INFOCROP rice model for each observation site in the eastern India under the three GCM scenarios
Sites

Bhubaneswar
Chinsurah
Cuttack
Faizabad
Jabalpur
Jorhat
Kalyani
Pusa
Raipur
Ranchi
Average change (%)
a

Rice yield
(t/ha)

GFDL
Predicted
change (%)

Predicted
yield (t/ha)

GISS
Predicted
change (%)

Predicted
yield (t/ha)

UKMO
Predicted
change (%)

Predicted
yield (t/ha)

4.46
5.18
4.93
4.72
7.54
3.83
3.55
3.82
3.75
4.5
4.63 

23.87 
7.03 
25.44 
13.55 
10.7
13.51 
8.73 
3.74 
1.71 
8.89 
9.02

3.40
4.82
3.68
4.08
6.73
4.35
3.24
3.68
3.69
4.10
4.18 

27.45 
7.38 
27.67 
17.65 
14.04
12.32 
11.65 
4.35 
5.11 
12.01 
11.50

3.24
4.80
3.57
3.89
6.48
4.30
3.14
3.65
3.56
3.96
4.06 

37.22 
8.11 
40.87 
28.34 
25.66
7.55 
22.38 
5.26 
18.01 
35.15 
21.35

2.80
4.76
2.92
3.38
5.61
4.12
2.76
3.62
3.07
2.92
3.60

Predicted rice yield is adjusted by the simulated changes in the experimental rice yield obtained.

Table 5b
Estimated changes in rice yield predicteda by the ORYZA1 model for each observation site in the eastern India under the three GCM scenarios
Sites

Bhubaneswar
Chinsurah
Cuttack
Faizabad
Jabalpur
Jorhat
Kalyani
Pusa
Raipur
Ranchi
Average change (%)
a

Rice yield
(t/ha)

GFDL
Predicted
change (%)

Predicted
yield (t/ha)

GISS
Predicted
change (%)

Predicted
yield (t/ha)

Predicted
change (%)

Predicted
yield (t/ha)

4.46
5.18
4.93
4.72
7.54
3.83
3.55
3.82
3.75
4.50
4.63 

17.33 
8.03 
19.67 
9.02 
11.05
12.13 
7.75 
4.93 
2.79 
7.87 
7.63

3.69
4.76
3.96
4.29
6.71
4.29
3.27
3.63
3.65
4.15
4.24 

20.36 
8.72 
20.32 
11.27 
14.08
12.64 
9.76 
6.31 
5.22 
10.35 
9.38

3.55
4.73
3.93
4.19
6.48
4.31
3.20
3.58
3.55
4.03
4.16 

27.53 
9.59 
30.75 
18.82 
21.05
8.31 
16.51 
6.58 
10.09 
25.98 
15.86

3.23
4.68
3.41
3.83
5.95
4.15
2.96
3.57
3.37
3.33
3.85

Predicted rice yield is adjusted by the simulated changes in the experimental rice yield obtained.

UKMO

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P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

suggested a significant decrease in rice yield in Cuttack,
while an increase in Jorhat under all three GCM scenarios.
The changes in yield components for both the sites are
provided in Table 6. In general, the number of spikelets
formed at both the locations was greater under the changed
climates. However, there was higher spikelet sterility at
Cuttack (>60%) than at Jorhat (13.25%). In addition, the
harvest indices at Cuttack decreased by about 25%, largely
due to the decreases in grain yield caused by supra-optimal
temperatures. On the contrary, there were only small
differences in harvest indices at Jorhat (Table 6).

Fig. 2. Changes in monthly average temperature (8C) obtained from the
daily weather data of each site during the main rice season (kharif) at
different sites in eastern India.

3.4. Methods of adaptation for higher rice yield under
climate change scenarios

3.3. Yield components at two contrasting sites

3.4.1. Adjustments in sowing date
Previous studies had suggested that adjusting sowing
dates might be a simple and powerful tool for mitigating the
effects of a potential global warming (Baker and Allen,
1993). The potential outcomes by adjusting the sowing time
in two sites (Cuttack and Jorhat) were examined by
simulating the crop growth under different climate change
scenarios. Under the GCMs scenarios, temperature at the
time of flowering in the main season was found to be high,
and there were considerable variations when simulated for
different climate change scenarios under different sowing
dates. Among the different sowing dates tested, the sowing
on July 15 at Cuttack led to the yield changes of +6.6, +4.1
and 9.8%, respectively, under the GFDL, GISS and
UKMO model scenarios (Fig. 3). Interestingly, the sowing
on July 1 at Jorhat resulted in yield increases of +27.1, +24.3
and +13.4%, respectively, for the corresponding scenarios
(Fig. 4). Any further delay in sowing at both the sites, which
had different dates for the maximum response, was not
beneficial in terms of crop yield (Figs. 3 and 4).

Among the different sites included for the present study,
two sites such as Cuttack, which showed the maximum yield
decrease, and Jorhat, which had the maximum increase in
yield, were selected to examine their responses under
different scenarios in detail. Interestingly, the results

3.4.2. Increased tolerance of spikelet fertility to
temperature
The hypothesis that high temperature induces spikelet
injury was evaluated by enhancing the tolerance level in
the crop models. The equation used in the ORYZA1 model

varied significantly: Cuttack (27.45%), Bhubaneswar
(25.63%), Ranchi (16.71%), Faizabad (16.44%),
Jabalpur (16.10%), Kalyani (12.80%), Chinsurah
(8.14%), Raipur (7.16%) and Pusa (5.20%). On the
contrary, there were increases in yield for Jorhat (11.08%).
Variation in average temperature during the normal growing
season is shown in Fig. 2. All the sites had higher maximum
average temperature at the time of sowing, followed by a
decrease in average temperature during tillering stage and an
increase during panicle initiation and flowering. The effect
of high temperature on spikelet sterility seems to be limited
to time of flowering and grain filling. During the flowering
stage, the maximum temperature was about 34 8C in Cuttack
and Bhubaneswar, around 33 8C in Jabalpur, 32 8C in
Faizabad, Kalyani and Chinsurah, 30 8C in Raipur and Pusa,
and 28 8C in Jorhat, which was the least of all.

Table 6
Simulated yield and yield components for the current climate and the climate change scenarios of the three GCMs at Cuttack and Jorhat
Yield
(kg ha1)

No. of spikelets
(m2)

No. of grains
(m2)

Filled grain
fraction (%)

1000 grain
weight (g)

Crop duration
(days)

Total dry
matter (kg ha1)

Harvest
index (%)

(i) Cuttack
Present
GFDL
GISS
UKMO

4930
3683
3572
2924

29,984
55,010
50,104
56,637

26,779
19,242
18,624
14,850

89.31
35.21
38.12
26.48

18.41
19.14
19.18
19.69

110
103
103
102

10564
12191
12653
11734

46.67
30.21
28.23
24.92

(ii) Jorhat
Present
GFDL
GISS
UKMO

3835
4357
4398
4197

21,636
25,945
25,900
25,702

20,640
22,401
22,600
22,148

95.40
87.12
87.53
86.75

18.58
19.45
19.46
18.95

120
105
105
103

8351
9318
9312
9493

45.92
46.76
47.23
44.21

P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

Fig. 3. Changes in yield (%) under the GCMs scenarios of the rice variety
IR 36 under different sowing dates grown during the kharif season at
Cuttack.

to describe the response of spikelet fertility to temperature
is

100
½1 þ

e0:853ðT max T mp Þ 

where d is the fertility percentage, Tmax the average daily
maximum temperature (8C) during the flowering period and
Tmp the average daily maximum temperature (8C) at which
50% of the spikelets are fertile. For the indica variety, Tmp
had a value of 36.5. To simulate the possible effect of an
increase in tolerance of spikelet to high temperatures, it was
assumed that this response was shifted by 2 8C by increasing
the value of Tmp to 38.5 8C. This adaptation in the spikelet
trait was examined in Cuttack site. With the available
weather data for this site, and with a constant sowing date
of June 15, a comparative study using the ORYZA1 model
was made for the current climate and other GCM scenarios
as obtained by the GFDL, GISS and UKMO (Fig. 5). Under
the GCMs scenarios, temperature at the time of flowering for
the main season was already high. Without any temperature

Fig. 4. Changes in yield (%) under the GCMs scenarios of the rice variety
IR 36 under different sowing dates grown during the kharif season at Jorhat.

239

Fig. 5. Changes in yield (%) under the GCMs scenarios of the rice variety
IR 36 and IR 36 with improved temperature tolerance grown during the
kharif season at Cuttack.

tolerance of the variety by not adjusting the value of Tmp,
large decreases in yield due to spikelet sterility were predicted. But with the adaptation of variety by improved
temperature tolerance of the spikelet, the yield increased
higher than that of the current scenario level, at about +10.7,
+13.6 and 8.4, respectively, under the GFDL, GISS and
UKMO model scenarios (Fig. 5).

4. Discussion
Both the crop simulation models predicted that any
increase in temperature at all the CO2 levels tested would
cause declines in yields but an increase in CO2 level at each
temperature increment would increase yields. These results
corroborated with that of Bachelet et al. (1995). Summarizing the data from several experimental studies on different
agricultural crops, Kimbal et al. (2002) found a 30%
increase in growth rate with a doubling of CO2 levels, which
was midway between the predicted values of the two models
in the present study. Nevertheless, the experimental findings
from the growth chamber studies (Baker et al., 1992a,b)
showed a 32% increase in rice grain yield due to doubling of
the CO2 concentration from 330 to 660 mmol CO2 mol1 air
(ppm). The increased growth response with increasing CO2
concentration was attributed to greater tillering and more
grain-bearing panicles. The net assimilation rate and canopy
net photosynthesis also increased with increasing CO2
concentration. The elevated CO2 concentration was found to
accelerate the development but shorten the total growth
duration of rice.
There are many indirect influences of elevated CO2 on
rice growth and development. When photosynthesis is
enhanced by increased CO2, the C/N ratio also increases in
the plants, which can reduce the nutritional quality of leaves
and increase feeding by the herbivorous insects (Johnson
and Lincoln, 1990). There can be considerable changes in
the nutrient-cycling processes in soils also (Strain, 1985).
Since the current crop growth simulation models have not

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P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

taken these factors into account, there are still limitations on
their predictive value.
The average yield changes of 8.23 and 7.31% by
ORYZA and INFOCROP, respectively, due to the effect of
temperature when simulated on per degree Celsius basis,
were comparable with that of 10% measured in the
controlled environment experiments (Baker and Allen,
1993). The low responses at 400 ppm and 1 8C in both the
models ORYZA (0.93%) and INFOCROP (2.90%) clearly
show the positive effects of temperature increase, simulated
by step-wise 1 8C increase with corresponding rise in CO2 to
400 ppm from the present ambient condition. However, the
CO2 concentration of 700 ppm resulted in increases of about
30.73 and 56.37% by ORYZA and INFOCROP, respectively.
In the light of recent experimental evidence (Kim et al.,
2003), these values appeared to be very high, probably
because the simulation models predicted the crop yield
mathematically from either RUE or net photosynthesis. In
ORYZA1, the hyperbolic relationship between the maximum rate of leaf photosynthesis at 1 g N/m2, and the
external CO2 concentration during rice growth has been
used. The rate of photosynthesis increased from
34 kg CO2 ha1 h1 at the 350 ppm CO2 concentration to
47 kg CO2 ha1 h1 at the 700 ppm CO2 concentration. The
CO2 fertilization factor is applied in INFOCROP to reflect
the direct physiological stimulation by elevated CO2
concentration. When compared with the results from the
FACE experiments (Kim et al., 2003), the fertilization
effects used in these two models are probably overestimated.
When simulated for the climate change scenarios, the
ORYZA model predicted changes of 7.63, 9.38 and 
15.86% for the GDFL, GISS and UKMO scenarios,
respectively, and INFOCROP predicted changes of 9.02, 
11.30 and 21.35%, respectively (Tables 5b and 5a). The
main cause for the differences in the predictions of the two
crop models was the way in which the leaf area development
and crop growth rate were calculated, and in the routines
describing phenological events in the crop. In ORYZA1, the
leaf area is calculated from leaf dry matter using the Specific
Leaf Area (SLA). LAI is simulated in the subroutine
SUBLAI. For a closed canopy, the LAI is calculated from
the leaf dry weight using SLA. When the canopy is not
closed, the plants grow exponentially as a function of the
temperature sum. The temperature sum is calculated using
the same procedure used to calculate the heat units for the
phenological development. The relative death rate of leaves
is applied to the leaf weight to calculate the weight loss of
leaves. The reduction in leaf area is calculated from the loss
of leaf weight using SLA (Kropff et al., 1994). In
INFOCROP, the Leaf Area Index changes proportionally
with Leaf Area Growth Rate (GLAI); its value is obtained by
multiplying the Increment in Leaf Weight (RWLVG) by the
SLA. The Net Leaf Area Growth Rate (RLAI) was
calculated based on the Initial Leaf Area Index (LAII),
GLAI, death rate of LAI (DLAI) and net loss of LAI due to
transplanting (PLTR) (Aggarwal et al., 2004).

In both the models, the phenological phases are
characterized by the thermal time and day length. In the
ORYZA1 model, the phenological development of the rice
crop is divided into four main phases, namely Basic
Vegetative Phase (BVP), Photoperiod-Sensitive Phase
(PSP), Panicle Formation Phase (PFP) and Grain Filling
Phase (GFP) (Kropff et al., 1994). In INFOCROP model, the
phenological development is divided into three main phases,
namely sowing to seedling emergence, seedling emergence
to anthesis and storage organ filling phase. The seedling
emergence to anthesis phase is further subdivided into three
major sub-phases depending on the environmental factors
affecting them and the organs formed, namely basic juvenile
phase, PSP and storage organ formation phase (Aggarwal
et al., 2004, 2006).
For each crop model, the GFDL scenario was the most
benign and the UKMO the most severe, corresponding to
the severity of temperature increases predicted by each GCM.
The predictions across both crop models and the three GCM
scenarios indicated a 12.45% decline in the overall regional
rice yield. Averaged across all three GCM scenarios, the mean
change in yield predicted by INFOCROP to be 13.95% and
by ORYZA to be 10.96%. Nevertheless, these values were
lesser than the average values for the scenarios in which
temperature and CO2 were varied at the fixed increments,
independently or in combination, above the current temperature for each site. It is likely that the GCM scenarios have
appropriate temperature corrections associated with the
elevated CO2 concentration, resulting in a better predictive
value compared to that of the scenarios with arbitrary
combinations of elevated CO2 and temperatures.
Among the different sites tested, both the models predicted the maximum loss in yield at Cuttack (27.45%),
while the maximum gain in yield was at Jorhat (+11.08%).
These differences in yield predictions were mainly due to the
rice spikelet sterility at high temperature. The temperature at
the time of flowering affects the spikelet fertility and hence
the yield (Krishnan and Surya Rao, 2005). The rice-growing
sites such as Cuttack and Bhubaneswar (hot, moist
subhumid climate type) had high maximum temperature
of about 34 8C and minimum temperature of 25 8C during
the flowering period. Other sites such as Jabalpur, Faizabad
and Ranchi (hot dry and moist subhumid type) had a high
maximum temperature of about 31 8C and the minimum
temperature of about 21 8C, which were lower than that of
Bhubaneswar and Cuttack. The low minimum temperature
probably helps to reduce the respiration at night. Likewise,
the rice-growing sites such as Kalyani, Pusa, Raipur and
Chinsurah (hot subhumid type) had a maximum temperature
of 30 8C and a minimum temperature of 20 8C during the
flowering period. But Jorhat (warm moist perhumid type)
had the maximum temperature of about 28 8C and a
minimum temperature of 19 8C only, which probably
contributed to the benefits from the predicted effects of
climate change scenarios. The predicted declines in the
overall rice yield by both cop models for the GFDL, GISS

P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

and UKMO scenarios showed the need to increase the rice
production than what was achieved at present. The
differences in the yield predicted due to different scenarios
were largely due to the differences in the temperature at the
time of flowering (Fig. 2).
Adjustment of management practices may help to offset
any detrimental effects of climate change on rice production.
Probably the easiest adaptation is to adjust the sowing dates.
Adjustment of sowing dates options was explored to
investigate a suitable agronomic option for adaptation
under the future climate change scenarios. In this way, for
the Cuttack site, the average yield changes of +6.6, +4.1 and 
9.8% were predicted during July 15 sowing for the GFDL,
GISS and UKMO scenarios, respectively, considerably
higher than those of 19.67, 20.32 and 30.75% observed
during June 15 sowing for the GFDL, GISS and UKMO
scenarios, respectively. Likewise, the Jorhat site showed
+27.1, +24.3 and +13.4% changes for the sowing on July 1
under the GFDL, GISS and UKMO model scenarios,
respectively; these changes were considerably higher than
those of +12.13, +12.64 and +8.31% for the June 15 sowing
for the corresponding scenarios. Further postponement in
sowing did not improve the grain yield, probably due to low
incident solar radiation and temperature.
There were striking differences in the predicted yield
changes among the three scenarios. The large differences
were mainly due to the sensitivity of spikelet sterility to
temperature. Even a small difference of just 1 8C could result
in a large yield decrease due to lower number of grains being
formed (Sheehy et al., 2006). This was illustrated in the two
examples used in the present study (Table 6). This was further
justified by the current changes in the temperature at different
sites during the main cropping season (Fig. 2). The imposed
climate change scenarios further enhanced this temperature
effect. The increased spikelet sterility at Cuttack was mainly
due to the predicted increment in temperature above the
already high daily maximum temperature (34 8C) (Prasad
et al., 2006; Krishnan and Surya Rao, 2005) under the climate
change scenarios reaching levels (38 8C) where spikelet
damage was considerable. Although the predicted temperature increments were similar to Cuttack, the lower average
temperature (28 8C) at Jorhat was well below the level (30 8C)
at which spikelet fertility is affected. At both the locations, the
number of spikelets formed was greater under the changed
climates. This could be due to the enhanced growth rate of the
crop between panicle initiation and flowering as the
consequences of fertilizing effect of higher CO2 level.
The limitation on yields imposed by the increased
spikelet sterility can be largely overcome by the selection of
genotypes that possess a higher potential of spikelet fertility
at high temperatures. The fertilizing effect of increased
atmospheric CO2 level is then likely to offset the changes in
crop development rate brought about by the increased
temperatures, so that significant yield increases may also be
obtained (Horie et al., 2000). Thus, the sensitivity of spikelet
sterility to temperature is a factor that must be taken into

241

account while evaluating the model predictions about the
effect of climate change on rice production. Further studies
on the adjustments to the management practices may help to
offset any detrimental effects of climate change on rice
production.
Some considerations are necessary, when interpreting
results from the scenarios predicted by the GCMs. The most
significant limitations are their poor resolution, inadequate
coupling of atmospheric and oceanic processes, poor
simulation of cloud processes and inadequate representation
of the biosphere and its feedbacks. The poor resolution is
likely to be significant in northeastern parts of India where
the relief is varied and local climate may be quite different
from the average across the area used by a GCM. Most
GCMs have difficulty in even describing the current climate
adequately (Bachelet et al., 1995). The current GCMs are
able to predict neither the changes in the variability of the
weather nor the frequency of catastrophic events such as
hurricanes, floods or even the intensity of monsoons, all of
which can be important in determining crop yields as the
average climatic data. It seems, therefore, that GCMs can at
best be used to suggest the likely direction and rate of change
of future climates.
According to Long et al. (2005), fertilization effect of
[CO2] has probably been overestimated, while omission of
[O3] effects from most models could have led to a 20%
overestimation of crop production in the Northern Hemisphere. Database of chamber studies are the mechanistic
basis for crop yield models. Hence, these models overestimate the yield gain due to elevated [CO2] compared to
those observed under fully open-air condition (FACE)
experiments in the field. The current FACE experiments are,
however, not adequate enough to reparameterize the existing
models (Long et al., 2005). In a recent study, Bannyayan
et al. (2005) evaluated ORYZA 2000 (Bouman and Van
Laar, 2006) against the observed growth and yield of rice in a
3-year field experiment in Japan where rice plants were
subjected to the elevated CO2 with FACE under varying N
fertilizer rates. The simulation results showed that the model
overestimated the increases in green leaf area indices due to
the elevated CO2 concentration but the enhancement of total
biomass was only a minor overestimation. While the model
was successful in simulating the increase in rice yield due to
the CO2 enrichment, it failed to reproduce the observe
interaction with N in the rice yield response to elevated CO2
concentration. Thus, the lack of complete understanding of
the effects and the potential interactions of environment
variables on plant processes preludes the definitive
predictions of the effects of global climate change.
Despite the limitations imposed by the assumptions made
in both the GCM and the crop simulation models, the current
study provides significant progress in our understanding of
how future climates are likely to affect rice production in the
eastern India. The use of simulation models to predict the
likely effects of climate change on crop production is an
evolving process. Our study in view of the findings of the

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P. Krishnan et al. / Agriculture, Ecosystems and Environment 122 (2007) 233–242

recent FACE studies clearly shows the need for modification
of the existing models. Other levels of production such as the
influences of water, nutrients and pests, and diseases due to
climate change are to be included in the refined models.
Some of these limitations in the use of present models can be
addressed so that increasingly more accurate predictions can
be made in future.

Acknowledgement
Our acknowledgements are to the collaborating members
from Orissa University of Agriculture & Technology
(OUAT), Bhubaneswar; Rice research Station (RRS),
Chinsurah; Narendra Deva University of Agriculture &
Technology (NDUAT), Faizabad; Jawaharlal Nehru Krishi
Vishwa Vidyalaaya (JNKVV), Jabalpur; Assam Agricultural
University (AAU), Jorhat; Bidan Chandra Krishi Viswa
Vidyalaya (BCKV), Kalyani, Rajendra Agricultural University (RAU), Pusa; Indira Gandhi Agricultural University
(IGAU), Raipur; Birsa Agricultural University (BAU),
Ranchi; for providing the crop parameter and weather data
under the NATP RRPS 25 and Indian Meteorological
Department (IMD), Pune, for weather data.

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