AGEE 3025 1–10

12 Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx

www.elsevier.com/locate/agee

4 Impact of elevated CO2 and temperature on rice yield and methods

F
of adaptation as evaluated by crop simulation studies

OO
5
P. Krishnan a,*, D.K. Swain b, S. Chandrabaskar b, S.K. Nayak a, R.N. Dash b
6
a
7 Division of Biochemistry, Plant Physiology and Environmental Sciences, Central Rice Research Institute, Cuttack 753006, India
b
8 Division of Soil Science and Microbiology, Central Rice Research Institute, Cuttack 753006, India

9 Received 7 February 2006; received in revised form 20 December 2006; accepted 19 January 2007

PR
10
11
12

13
Abstract
14
15 Impact of elevated CO2 and temperature on rice yield in eastern India was simulated by using the ORYZA1 and the INFOCROP rice
16
17
ED
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
18 1 8C increase in temperature, ORYZA1 and INFOCROP rice models predicted average yield changes of 7.20 and 6.66%, respectively, at
19 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%
20 by ORYZA1 and 56.37% by INFOCROP rice. When temperature was increased by about +4 8C above the ambient level, the differences in the
21 responses by the two models became remarkably small. For the GDFL, GISS and UKMO scenarios, ORYZA1 predicted the yield changes of
CT

22 7.63, 9.38 and 15.86%, respectively, while INFOCROP predicted changes of 9.02, 11.30 and 21.35%. There were considerable
23 differences in the yield predictions for individual sites, with declining trend for Cuttack and Bhubaneswar but an increasing trend for Jorhat.
24 These differences in yield predictions were mainly attributed to the sterility of rice spikelets at higher temperatures. Results suggest that the
25 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
26 of genotypes that possess higher fertility of spikelets at high temperatures.
E

27 # 2007 Published by Elsevier B.V.

28 Keywords: Climate change; CO2; INFOCROP; ORYZA; Oryza sativa L.; Simulation; Temperature; Yield
RR

29

30 33
1. Introduction their potential threat to rice productivity and the associated 34
31 impact on the socioeconomic structure of many rice-growing 35
CO

32 The climatic variability and the predicted climatic changes
33 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;
UN
countries. Among the global atmospheric changes, the incre- 36
asing concentrations of greenhouse gases such as CO2 may 37
have significant effect on rice productivity due to increase in 38
both the average surface temperature and the amount of CO2 39
available for photosynthesis (Aggarwal, 2003). In the absence 40
of temperature increase, many studies have shown that the net

41
GFP, Grain Filling Phase; GISS, Goddard Institute of Space Studies; GLAI,
Leaf Area Growth Rate; IPCC, Intergovernmental Panel on Climate
effect of doubling of CO2 was increase in the yield of rice 42
Change; LAI, Leaf Area Index; NATP, National Agricultural Technology (Kim et al., 2003). It becomes necessary to assess the effects of 43
Project; PFP, Panicle Formation Phase; PLTR, Net loss of LAI due to potential interactive changes of CO2 and temperature in order 44
transplanting; PSP, Photoperiod-Sensitive Phase; RLAI, Net Leaf Area to determine the future agricultural strategies that would 45
Growth Rate; RUE, Radiation Use Efficiency; RWLVG, Increment in Leaf maintain higher rice productivity. 46
Weight; SLA, Specific Leaf Area; SUBLAI, LAI is simulated in the
subroutine SUBLAI; UKMO, United Kingdom Meteorological Office
The simulations by different models and many field 47
* Corresponding author. experiments have shown the potential impact of climatic 48
E-mail address: prameelakrishnan@yahoo.com (P. Krishnan). change and the variability in rice productivity (Baker et al., 49

0167-8809/$ – see front matter # 2007 Published by Elsevier B.V.

doi:10.1016/j.agee.2007.01.019

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

2 P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx

49
50 1992; Peng et al., 2004; Kim et al., 2003). The modeling
51 studies from Bangladesh (Karim et al., 1994), Japan (Horie
52 et al., 2000), China (Bachelet et al., 1995) and India (Mall
53 and Aggarwal, 2002) reported the country-wise variations in
54 rice production, anticipated due to the climatic changes. The
55 simulated yields increased when temperature increases were
56 small, but declined when the decadal temperature increase
57 was more than 0.8 8C, with the greatest decline in crop

F
58 yields occurring between the latitudes of 108 and 358N.
59 Similar results were obtained by Penning de Vries (1993).

OO
60 Many uncertainties exist in modeling studies, partly due
61 to the quality of the predictions by the models, from the use
62 of limited sites for which historical weather data are
63 available, due to the quality of the crop simulation models,
Fig. 1. Comparison of yield simulated by ORYZA1 and INFOCROP rice.
64 especially when applied under the rain-fed conditions
65 (Bachelet et al., 1995), and due to the quality of the climate 102

PR
66 models used to predict future weather scenarios. These matter production as a function of light, CO2 and tempe- 103
67 uncertainties may be reduced only when a large number of rature by considering photosynthetic processes at the leaf 104
68 scenarios for different locations are compared and evaluated. level and integrating these over the canopy to obtain crop- 105
69 In order to overcome the predicted limitations for rice level values. Respiration is also modeled explicitly as a 106
70 production in the future, there is also a need to identify and function of temperature and partitioning of dry matter is 107
71 evaluate the suitable agronomic practices such as altered ED according to phenology-dependent functions. Thus, ORYZA 108
72 sowing date and selection of improved varieties with calculates dry matter production as a function of gross 109
73 increased spikelet fertility at high temperature and other canopy photosynthesis, depending on the detailed calcula- 110
74 useful traits. tions of the distribution of light within the canopies, 111
75 Attempts have been made earlier to assess the general the radiation absorbed by the canopy and photosynthesis 112
76 effects of global environmental changes on rice yield using light response cure of leaves. Growth and maintenance 113
CT

77 simulation models, but little attention has been given to the respiration are calculated as a function of tissue N content, 114
78 potential interactive effects between high temperature and temperature and crop-specific coefficients. This methodol- 115
79 increasing CO2 levels. Lal et al. (1998) used the CERES ogy, although yields very accurate results, poses practical 116
80 rice and predicted a 20% decline in rice yields in the difficulties because of its requirement for detailed and 117
81 northwestern India due to elevated CO2 and temperature. careful measurements (Kropff et al., 1994). The INFOCROP 118
E

82 Eastern India accounts for about 63% (26.5 million ha) of model, however, uses a simpler radiation use efficiency 119
83 the total rice-growing area in India. The rice ecosystems in (RUE) relationship between intercepted solar radiation and 120
RR

84 these regions show characteristic differences with respect to growth, in which respiration is implicit. CO2 effects are 121
85 the environmental factors as well as the cultural practices. accounted for by using a curvilinear function relating RUE 122
86 There is an urgent need to characterize the impact of future to CO2 concentration. Temperature is assumed not to affect 123
87 climatic changes on rice yield in these regions for sustaining RUE. More or less similar results can be obtained under 124
88 the productivity. The present paper discusses the outcomes normal radiation situations by calculating the net dry matter 125
CO

89 of two rice growth simulation models, ORYZA and production as a function of RUE. Pre-determined values of 126
90 INFOCROP rice, when applied to different rice-growing RUE were input in the model as a function of crop/cultivar 127
91 sites in the eastern India for studying the potential interactive (RUEMAX), and RUE was further modified by the develop- 128
92 effects between high temperature and increasing CO2 levels ment stage (Aggarwal et al., 2004). 129
93 and for the different thermal climate change scenarios. The rice-growing regions included for the present study 130
lie in eastern India, and these sites (Bhubaneswar, Chin- 131
UN

surah, Cuttack, Faizabad, Jabalpur, Jorhat, Kalyani, Pusa, 132

2. Materials and methods Raipur and Ranchi), which are geographically apart at varied 133
94 altitudes ranging from 7.8 to 86.56 m above mean sea level, 134
95 Two popular models of rice growth ORYZA1 (Kropff show characteristic features with respect to the weather and 135
96 et al., 1994) and INFOCROP rice (Aggarwal et al., 2006) are crop factors. The crop parameters included in the study were 136
97 used in this study. Prior to their use, both were evaluated obtained from the field experiments conducted at these sites 137
98 and compared (Fig. 1) and then these crop models were under National Agricultural Technology Project (NATP) 138
99 calibrated for the indica variety IR 36 at all sites. In general, RRPS 25 during 2001–2004 (Table 1). 139
100 the two models differ in the manner in which dry matter The input parameters used for the two models are given in 140
101 production and partition, leaf area development, and pheno- Table 2a (parameters such as varietal data, soil data and 141
102 logical development are calculated. ORYZA1 calculates dry weather data), Table 2b (physical and chemical properties of 142

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx 3

146
Table 1 were made for the main rice-growing season for each of 147
Sites and institutes where the field observation and weather data were these sites. Generally, the dates of sowing and transplanting 148
collected
were supplied along with the weather data. 149
Sites Institutes
Bhubaneswar Orissa University of Agriculture & Technology (OUAT) 2.1. Climate change scenarios
Chinsurah Rice Research Station (RRS)
150
Cuttack Central Rice Research Institute (CRRI)
Faizabad Narendra Deva University of Agriculture & For each of these rice-growing regions, the potential yield 151
Technology (NDUAT) of rice was simulated under 30 different combinations of 152

F
Jabalpur Jawaharlal Nehru Krishi Vishwa Vidyalaya (JNKVV) CO2 and temperature, including with the ‘fixed increment’ 153
Jorhat Assam Agricultural University (AAU) changes in CO2 (380, 400, 500, 600 and 700 ppm) and 154

OO
Kalyani Bidan Chandra Krishi Viswa Vidyalaya (BCKV)
temperature (ambient, +1, +2, +3, +4 and +5 8C) individu- 155
Pusa Rajendra Agricultural University (RAU)
Raipur Indira Gandhi Agricultural University (IGAU) ally, and with all the combinations of these levels of CO2 and 156
Ranchi Birsa Agricultural University (BAU) temperature. The actual daily weather data, collected for 157
Indian Meteorological Department (IMD), three consecutive years from each of these sites were used 158
Pune (weather data) for simulation. For temperature changes, the daily weather 159
142 data from each site were exported to Excel spread sheets and 160

PR
143 soils) and Table 2c (geographical information). The weather then the daily maximum and minimum temperatures were 161
144 data containing measurements of sunshine hours, tempera- increased by 1–5 8C individually. Later, the modified 162
145 ture and rainfall from all these sites were used as the baseline weather data were used as the inputs for ORYZA1. In case 163
146 data for input into these crop models. Later, the simulations of INFOCROP, for a given daily weather data, the climate 164

Table 2a ED
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 1. Maximum and minimum
temperatures
2. Thermal time for sowing to germination 2. Soil texture 2. Relative humidity
CT
3. Base temperature for germination to 50% flowering 3. pH of soil 3. Sun shine hours
4. Thermal time for germination to 50% flowering 4. EC 4. Precipitation
5. Base temperature for 50% flowering to physiological maturity 5. Sand %
6. Thermal time for 50% flowering to physiological maturity 6. Silt %
7. Optimal temperature 7. Clay %
8. Maximum temperature 8. Saturation fraction
E

9. Sensitivity to photoperiod 9. Field capacity

10. Relative growth rate of leaf area 10. Wilting point
11. Specific Leaf Area 11. Saturated hydraulic conductivity
RR

12. Extinction coefficient of leaves at flowering 12. Bulk density

13. Radiation Use Efficiency 13. Organic carbon
14. Root growth rate
15. Index of greenness of leaves
16. Sensitivity of crop to flooding
17. Index of N fixation
CO

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
UN

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 Sandy clay Sandy clay Loam Loam Sandy clay Silt Clayey Sandy clay Sandy
loam loam loam loam loam
B.D.a (g/cc) 1.35 1.45 1.3 1.41 1.3 1.23 1.34 1.48 1.4
O.C.b (%) 0.73 0.51 0.3 0.68 0.57 0.54 0.69 0.67 0.47
Total N (%) 0.07 0.04 – 0.07
Available N (kg/ha) 128.4 355 258.6 – 267 265.8
a
B.D., bulk density.
b
O.C., organic carbon.

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

4 P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx

Table 2c
Model inputs of geographical information of different locations
Soil properties Bhubaneswar Kalyani Faizabad Ranchi Pusa Raipur Jabalpur Jorhat Cuttack
0 0 0 0 0 0 0 0
Latitude 20814 N 22857 N 22887 N 23817 N 2285 N 25859 N 23809 N 2688 N 208300 N
Longitude 858520 E 888210 E 88840 E 858190 E 868E 85840 E 798580 E 95850 E 868E
Altitude (m) 25.9 7.8 8.62 625 23 51.84 411 86.56 23

164 202
165 change scenarios were specified as +1 to +5 8C for relative increase predicted for each site was weighed by its 203

F
166 temperatures and the percent changes in precipitation. current production observed in the field trials of an earlier 204
167 The inbuilt weather generator of that model generated the study (Annual Report of NATP RRPS 25, 2002–2003). The 205

OO
168 modified daily weather data accordingly, and these data were differences in the production capacities among these sites 206
169 used as the daily weather inputs. The climate change were evaluated. 207
170 scenarios under different levels of CO2 were applied by
171 changing the ambient CO2 concentration parameter in these
172 two models. 3. Results
173 Besides, the outputs from the climate models were used 208

PR
174 in combination with these crop models. The coarse grid from 3.1. Effect of temperature and CO2 levels at fixed 209
175 each GCM was interpolated using a four point inverse- increments on yields
176 distance-squared algorithm to a 0.58 latitude  0.58 long- 210
177 itude grid using a raster-based Geographical Information At all the CO2 levels tested (380, 400, 500, 600 and 211
178 Systems (GIS) software package. Scenarios were produced 700 ppm), both the models predicted the declining yields of 212
179 by applying ratios of precipitation or differences in tempe- ED rice due to an increase in temperature. On the contrary, an 213
180 rature predicted for the 2  CO2 and 1  CO2 simulation to increase in CO2 level at any particular temperature increased 214
181 the baseline present climate data set for different sites the rice yields (Table 4). At the current level of CO2 215
182 (Mathews and Wassmann, 2003). The important features of (considered at 380 ppm), ORYZA predicted a mean change 216
183 three General Circulation Models (GCMs) used such as of 7.20% in yields for every 1 8C increase in temperature, 217
184 the General Fluid Dynamics Laboratory (GFDL) Model, while INFOCROP predicted 6.66%. But increasing CO2 218
CT

185 Goddard Institute of Space Studies (GISS) model and the concentration (700 ppm) resulted in increases of 30.37 and 219
186 United Kingdom Meteorological Office (UKMO) model are 56.37% in yield by ORYZA and INFOCROP, respectively. 220
187 provided in Table 3. However, with temperature increase of +4 8C above 221
188 The GCM scenarios were produced by applying the ratios ambient, the differences in the yield predictions by the 222
189 of precipitation or differences in temperature predicted for two models became remarkably small (Table 4). 223
E

190 the 1  CO2 (380 ppm) and 2  CO2 (760 ppm) simulations
191 to the baseline daily weather data set. The changes between 3.2. Effect of predicted GCM scenarios on rice yields
RR

192 1  CO2 and 2  CO2 conditions were representative of the 224

193 differences between the present and the future climate The predicted changes in overall production for each 225
194 scenarios following an equivalent doubling of CO2. site under different climate scenarios using the two crop 226
models are provided in Tables 5a and 5b. In general, the 227
2.2. Simulation ORYZA model suggested the decreases of 7.63, 9.38 228
CO

195 and 15.86% in yield for the GDFL, GISS and UKMO 229
196 For the simulation analysis, only runs that terminated scenarios, respectively. For the corresponding scenarios, 230
197 normally or by crop deaths as a result of temperature were INFOCROP indicated larger reductions at 9.02, 11.30 231
198 included. The relative yield changes under the scenarios, and 21.35%, respectively. When each site was analysed 232
199 referred to yields predicted for the current climate, were individually under three GISS and UKMO scenarios, almost 233
200 used in the analysis rather than absolute yields. To provide all the sites except one showed the declining trend in yields. 234
UN

201 an estimate of the overall effect of climate change on rice The decreases in the yield of rice, when the mean of the 235
202 production under the three GCMs scenarios, the average corresponding values in Tables 5a and 5b were considered, 236

Table 3
The major features of the General Circulation Models used in this study
GFDL GISS UKMO
Source laboratory Geophysical Fluid Dynamics Goddard Institute for United Kingdom
Laboratory Space Studies Meteorological Office
Base 1  CO2 (ppm) 380 380 380
Change in global temperature (8C) +4.0 +4.2 +5.2
Change in global precipitation (%) 8 11 15

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
AGEE 3025 1–10

P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx 5

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 0 2.68 18.04 28.15 36.36 42.49 20.39
400 5.23 2.90 12.03 23.21 33.05 40.32 18.12
500 25.92 13.90 3.60 12.32 25.08 35.11 4.85

F
600 40.61 29.39 16.74 1.04 15.80 29.17 7.14
700 56.37 41.21 27.96 9.85 7.67 23.15 17.43
Average 25.14 16.86 3.65 10.56 23.59 34.05 3.76

OO
ORYZA
380 0 4.27 15.32 26.78 29.45 38.96 19.13
400 2.78 0.93 8.44 19.74 23.67 33.65 13.22
500 18.57 11.73 0.24 11.75 15.43 26.57 3.87
600 26.8 18.54 5.03 6.86 10.87 21.84 1.80
700 30.73 23.77 8.67 3.68 7.32 19.04 5.52

PR
Average 16.27 10.14 1.96 13.76 17.35 28.01 5.78
a
Changes are averaged across all sites and at all the available years.
b
Temperature increments are above the current temperatures at each site.

Table 5a ED
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 Rice yield GFDL GISS UKMO
(t/ha) Predicted Predicted Predicted Predicted Predicted Predicted
change (%) yield (t/ha) change (%) yield (t/ha) change (%) yield (t/ha)
CT
Bhubaneswar 4.46 23.87 3.40 27.45 3.24 37.22 2.80
Chinsurah 5.18 7.03 4.82 7.38 4.80 8.11 4.76
Cuttack 4.93 25.44 3.68 27.67 3.57 40.87 2.92
Faizabad 4.72 13.55 4.08 17.65 3.89 28.34 3.38
Jabalpur 7.54 10.7 6.73 14.04 6.48 25.66 5.61
Jorhat 3.83 13.51 4.35 12.32 4.30 7.55 4.12
E

Kalyani 3.55 8.73 3.24 11.65 3.14 22.38 2.76

Pusa 3.82 3.74 3.68 4.35 3.65 5.26 3.62
Raipur 3.75 1.71 3.69 5.11 3.56 18.01 3.07
RR

Ranchi 4.5 8.89 4.10 12.01 3.96 35.15 2.92

Average change (%) 4.63 9.02 4.18 11.50 4.06 21.35 3.60
a
Predicted rice yield is adjusted by the simulated changes in the experimental rice yield obtained.
CO

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 Rice yield GFDL GISS UKMO
(t/ha) Predicted Predicted Predicted Predicted Predicted Predicted
change (%) yield (t/ha) change (%) yield (t/ha) change (%) yield (t/ha)
UN

Bhubaneswar 4.46 17.33 3.69 20.36 3.55 27.53 3.23

Chinsurah 5.18 8.03 4.76 8.72 4.73 9.59 4.68
Cuttack 4.93 19.67 3.96 20.32 3.93 30.75 3.41
Faizabad 4.72 9.02 4.29 11.27 4.19 18.82 3.83
Jabalpur 7.54 11.05 6.71 14.08 6.48 21.05 5.95
Jorhat 3.83 12.13 4.29 12.64 4.31 8.31 4.15
Kalyani 3.55 7.75 3.27 9.76 3.20 16.51 2.96
Pusa 3.82 4.93 3.63 6.31 3.58 6.58 3.57
Raipur 3.75 2.79 3.65 5.22 3.55 10.09 3.37
Ranchi 4.50 7.87 4.15 10.35 4.03 25.98 3.33
Average change (%) 4.63 7.63 4.24 9.38 4.16 15.86 3.85
a
Predicted rice yield is adjusted by the simulated changes in the experimental rice yield obtained.

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

6 P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx

258
suggested a significant decrease in rice yield in Cuttack, 259
while an increase in Jorhat under all three GCM scenarios. 260
The changes in yield components for both the sites are 261
provided in Table 6. In general, the number of spikelets 262
formed at both the locations was greater under the changed 263
climates. However, there was higher spikelet sterility at 264
Cuttack (>60%) than at Jorhat (13.25%). In addition, the 265
harvest indices at Cuttack decreased by about 25%, largely 266

F
due to the decreases in grain yield caused by supra-optimal 267
temperatures. On the contrary, there were only small 268

OO
differences in harvest indices at Jorhat (Table 6). 269

3.4. Methods of adaptation for higher rice yield under 270

Fig. 2. Changes in monthly average temperature (8C) obtained from the climate change scenarios
daily weather data of each site during the main rice season (kharif) at 271
different sites in eastern India.
3.4.1. Adjustments in sowing date 272

PR
236 Previous studies had suggested that adjusting sowing 273
237 varied significantly: Cuttack (27.45%), Bhubaneswar dates might be a simple and powerful tool for mitigating the 274
238 (25.63%), Ranchi (16.71%), Faizabad (16.44%), Jabal- effects of a potential global warming (Baker and Allen, 275
239 pur (16.10%), Kalyani (12.80%), Chinsurah (8.14%), 1993). The potential outcomes by adjusting the sowing 276
240 Raipur (7.16%) and Pusa (5.20%). On the contrary, there time in two sites (Cuttack and Jorhat) were examined by 277
241 were increases in yield for Jorhat (11.08%). Variation in ED simulating the crop growth under different climate change 278
242 average temperature during the normal growing season is scenarios. Under the GCMs scenarios, temperature at the 279
243 shown in Fig. 2. All the sites had higher maximum average time of flowering in the main season was found to be high, 280
244 temperature at the time of sowing, followed by a decrease in and there were considerable variations when simulated for 281
245 average temperature during tillering stage and an increase different climate change scenarios under different sowing 282
246 during panicle initiation and flowering. The effect of high dates. Among the different sowing dates tested, the sowing 283
CT

247 temperature on spikelet sterility seems to be limited to time of on July 15 at Cuttack led to the yield changes of +6.6, +4.1 284
248 flowering and grain filling. During the flowering stage, the and 9.8%, respectively, under the GFDL, GISS and 285
249 maximum temperature was about 34 8C in Cuttack and UKMO model scenarios (Fig. 3). Interestingly, the sowing 286
250 Bhubaneswar, around 33 8C in Jabalpur, 32 8C in Faizabad, on July 1 at Jorhat resulted in yield increases of +27.1, +24.3 287
251 Kalyani and Chinsurah, 30 8C in Raipur and Pusa, and 28 8C and +13.4%, respectively, for the corresponding scenarios 288
E

252 in Jorhat, which was the least of all. (Fig. 4). Any further delay in sowing at both the sites, which 289
had different dates for the maximum response, was not 290
RR

3.3. Yield components at two contrasting sites beneficial in terms of crop yield (Figs. 3 and 4). 291
253
254 Among the different sites included for the present study, 3.4.2. Increased tolerance of spikelet fertility to 292
255 two sites such as Cuttack, which showed the maximum temperature 293
256 yield decrease, and Jorhat, which had the maximum increase The hypothesis that high temperature induces spikelet 294
CO

257 in yield, were selected to examine their responses under injury was evaluated by enhancing the tolerance level in 295
258 different scenarios in detail. Interestingly, the results the crop models. The equation used in the ORYZA1 model 296

Table 6
Simulated yield and yield components for the current climate and the climate change scenarios of the three GCMs at Cuttack and Jorhat
UN

Yield No. of spikelets No. of grains Filled grain 1000 grain Crop duration Total dry Harvest
(kg ha1) (m2) (m2) fraction (%) weight (g) (days) matter (kg ha1) index (%)
(i) Cuttack
Present 4930 29,984 26,779 89.31 18.41 110 10564 46.67
GFDL 3683 55,010 19,242 35.21 19.14 103 12191 30.21
GISS 3572 50,104 18,624 38.12 19.18 103 12653 28.23
UKMO 2924 56,637 14,850 26.48 19.69 102 11734 24.92
(ii) Jorhat
Present 3835 21,636 20,640 95.40 18.58 120 8351 45.92
GFDL 4357 25,945 22,401 87.12 19.45 105 9318 46.76
GISS 4398 25,900 22,600 87.53 19.46 105 9312 47.23
UKMO 4197 25,702 22,148 86.75 18.95 103 9493 44.21

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx 7

F
OO
Fig. 5. Changes in yield (%) under the GCMs scenarios of the rice variety
Fig. 3. Changes in yield (%) under the GCMs scenarios of the rice variety IR 36 and IR 36 with improved temperature tolerance grown during the
IR 36 under different sowing dates grown during the kharif season at kharif season at Cuttack.
Cuttack.

PR
296 314
297 to describe the response of spikelet fertility to temperature tolerance of the variety by not adjusting the value of Tmp, 315
is large decreases in yield due to spikelet sterility were pre- 316
dicted. But with the adaptation of variety by improved 317
100 temperature tolerance of the spikelet, the yield increased 318
299
298
d¼
½1 þ e0:853ðT max T mp Þ  higher than that of the current scenario level, at about +10.7,
ED 319
300 +13.6 and 8.4, respectively, under the GFDL, GISS and 320
301 where d is the fertility percentage, Tmax the average daily UKMO model scenarios (Fig. 5). 321
302 maximum temperature (8C) during the flowering period and
303 Tmp the average daily maximum temperature (8C) at which
304 50% of the spikelets are fertile. For the indica variety, Tmp 4. Discussion
CT

305 had a value of 36.5. To simulate the possible effect of an 322

306 increase in tolerance of spikelet to high temperatures, it was Both the crop simulation models predicted that any 323
307 assumed that this response was shifted by 2 8C by increasing increase in temperature at all the CO2 levels tested would 324
308 the value of Tmp to 38.5 8C. This adaptation in the spikelet cause declines in yields but an increase in CO2 level at 325
309 trait was examined in Cuttack site. With the available each temperature increment would increase yields. These 326
E

310 weather data for this site, and with a constant sowing date results corroborated with that of Bachelet et al. (1995). 327
311 of June 15, a comparative study using the ORYZA1 model Summarizing the data from several experimental studies on 328
RR

312 was made for the current climate and other GCM scenarios different agricultural crops, Kimbal et al. (2002) found a 329
313 as obtained by the GFDL, GISS and UKMO (Fig. 5). Under 30% increase in growth rate with a doubling of CO2 levels, 330
314 the GCMs scenarios, temperature at the time of flowering for which was midway between the predicted values of the two 331
the main season was already high. Without any temperature models in the present study. Nevertheless, the experimental 332
findings from the growth chamber studies (Baker et al., 1990 333
CO

a,b) showed a 32% increase in rice grain yield due to 334

doubling of the CO2 concentration from 330 to 660 mmol - 335
CO2 mol1 air (ppm). The increased growth response with 336
increasing CO2 concentration was attributed to greater 337
tillering and more grain-bearing panicles. The net assimila- 338
tion rate and canopy net photosynthesis also increased with 339
UN

increasing CO2 concentration. The elevated CO2 concentra- 340

tion was found to accelerate the development but shorten the 341
total growth duration of rice. 342
There are many indirect influences of elevated CO2 343
on rice growth and development. When photosynthesis is 344
enhanced by increased CO2, the C/N ratio also increases in 345
the plants, which can reduce the nutritional quality of leaves 346
and increase feeding by the herbivorous insects (Johnson 347
and Lincon, 1990). There can be considerable changes in the 348
Fig. 4. Changes in yield (%) under the GCMs scenarios of the rice variety nutrient-cycling processes in soils also (Strain, 1985). Since 349
IR 36 under different sowing dates grown during the kharif season at Jorhat. the current crop growth simulation models have not taken 350

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

8 P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx

350 406
351 these factors into account, there are still limitations on their In both the models, the phenological phases are charac- 407
352 predictive value. terized by the thermal time and day length. In the ORYZA1 408
353 The average yield changes of 8.23 and 7.31% by model, the phenological development of the rice crop is 409
354 ORYZA and INFOCROP, respectively, due to the effect divided into four main phases, namely Basic Vegetative 410
355 of temperature when simulated on per degree Celsius Phase (BVP), Photoperiod-Sensitive Phase (PSP), Panicle 411
356 basis, were comparable with that of 10% measured in the Formation Phase (PFP) and Grain Filling Phase (GFP) 412
357 controlled environment experiments (Baker and Allen, (Kropff et al., 1994). In INFOCROP model, the phenological 413
358 1993). The low responses at 400 ppm and 1 8C in both the development is divided into three main phases, namely 414

F
359 models ORYZA (0.93%) and INFOCROP (2.90%) clearly sowing to seedling emergence, seedling emergence to 415
360 show the positive effects of temperature increase, simulated anthesis and storage organ filling phase. The seedling 416

OO
361 by step-wise 1 8C increase with corresponding rise in CO2 to emergence to anthesis phase is further subdivided into three 417
362 400 ppm from the present ambient condition. However, the major sub-phases depending on the environmental factors 418
363 CO2 concentration of 700 ppm resulted in increases of about affecting them and the organs formed, namely basic juvenile 419
364 30.73 and 56.37% by ORYZA and INFOCROP, respectively. phase, PSP and storage organ formation phase (Aggarwal 420
365 In the light of recent experimental evidence (Kim et al., et al., 2004, 2006). 421
366 2003), these values appeared to be very high, probably For each crop model, the GFDL scenario was the most 422

PR
367 because the simulation models predicted the crop yield benign and the UKMO the most severe, corresponding to the 423
368 mathematically from either RUE or net photosynthesis. In severity of temperature increases predicted by each GCM. 424
369 ORYZA1, the hyperbolic relationship between the max- The predictions across both crop models and the three 425
370 imum rate of leaf photosynthesis at 1 g N/m2, and the GCM scenarios indicated a 12.45% decline in the overall 426
371 external CO2 concentration during rice growth has been regional rice yield. Averaged across all three GCM scena- 427
372 used. The rate of photosynthesis increased from 34 kg ED rios, the mean change in yield predicted by INFOCROP to 428
373 CO2 ha1 h1 at the 350 ppm CO2 concentration to 47 kg be 13.95% and by ORYZA to be 10.96%. Nevertheless, 429
374 CO2 ha1 h1 at the 700 ppm CO2 concentration. The CO2 these values were lesser than the average values for the 430
375 fertilization factor is applied in INFOCROP to reflect the scenarios in which temperature and CO2 were varied at the 431
376 direct physiological stimulation by elevated CO2 concen- fixed increments, independently or in combination, above 432
377 tration. When compared with the results from the FACE the current temperature for each site. It is likely that the 433
CT

378 experiments (Kim et al., 2003), the fertilization effects used GCM scenarios have appropriate temperature corrections 434
379 in these two models are probably overestimated. associated with the elevated CO2 concentration, resulting in 435
380 When simulated for the climate change scenarios, the a better predictive value compared to that of the scenarios 436
381 ORYZA model predicted changes of 7.63, 9.38 and with arbitrary combinations of elevated CO2 and tempera- 437
382 15.86% for the GDFL, GISS and UKMO scenarios, tures. 438
E

383 respectively, and INFOCROP predicted changes of 9.02, Among the different sites tested, both the models pre- 439
384 11.30 and 21.35%, respectively (Tables 5b and 5a). The dicted the maximum loss in yield at Cuttack (27.45%), 440
RR

385 main cause for the differences in the predictions of the two while the maximum gain in yield was at Jorhat (+11.08%). 441
386 crop models was the way in which the leaf area development These differences in yield predictions were mainly due to the 442
387 and crop growth rate were calculated, and in the routines rice spikelet sterility at high temperature. The temperature at 443
388 describing phenological events in the crop. In ORYZA1, the the time of flowering affects the spikelet fertility and hence 444
389 leaf area is calculated from leaf dry matter using the Specific the yield (Krishnan and Surya Rao, 2005). The rice-growing 445
CO

390 Leaf Area (SLA). LAI is simulated in the subroutine sites such as Cuttack and Bhubaneswar (hot, moist sub- 446
391 SUBLAI. For a closed canopy, the LAI is calculated from humid climate type) had high maximum temperature of 447
392 the leaf dry weight using SLA. When the canopy is not about 34 8C and minimum temperature of 25 8C during the 448
393 closed, the plants grow exponentially as a function of the flowering period. Other sites such as Jabalpur, Faizabad and 449
394 temperature sum. The temperature sum is calculated using Ranchi (hot dry and moist subhumid type) had a high 450
395 the same procedure used to calculate the heat units for the maximum temperature of about 31 8C and the minimum 451
UN

396 phenological development. The relative death rate of leaves temperature of about 21 8C, which were lower than that of 452
397 is applied to the leaf weight to calculate the weight loss Bhubaneswar and Cuttack. The low minimum temperature 453
398 of leaves. The reduction in leaf area is calculated from the probably helps to reduce the respiration at night. Likewise, 454
399 loss of leaf weight using SLA (Kropff et al., 1994). In the rice-growing sites such as Kalyani, Pusa, Raipur and 455
400 INFOCROP, the Leaf Area Index changes proportionally Chinsurah (hot subhumid type) had a maximum temperature 456
401 with Leaf Area Growth Rate (GLAI); its value is obtained by of 30 8C and a minimum temperature of 20 8C during the 457
402 multiplying the Increment in Leaf Weight (RWLVG) by flowering period. But Jorhat (warm moist perhumid type) 458
403 the SLA. The Net Leaf Area Growth Rate (RLAI) was had the maximum temperature of about 28 8C and a 459
404 calculated based on the Initial Leaf Area Index (LAII), minimum temperature of 19 8C only, which probably contri- 460
405 GLAI, death rate of LAI (DLAI) and net loss of LAI due to buted to the benefits from the predicted effects of climate 461
406 transplanting (PLTR) (Aggarwal et al., 2004). change scenarios. The predicted declines in the overall rice 462

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx 9

462 518
463 yield by both cop models for the GFDL, GISS and UKMO temperatures, so that significant yield increases may also be 519
464 scenarios showed the need to increase the rice production obtained (Horie et al., 2000). Thus, the sensitivity of spikelet 520
465 than what was achieved at present. The differences in the sterility to temperature is a factor that must be taken into 521
466 yield predicted due to different scenarios were largely due to account while evaluating the model predictions about the 522
467 the differences in the temperature at the time of flowering effect of climate change on rice production. Further studies 523
468 (Fig. 2). on the adjustments to the management practices may help to 524
469 Adjustment of management practices may help to offset offset any detrimental effects of climate change on rice 525
470 any detrimental effects of climate change on rice production. production. 526

F
471 Probably the easiest adaptation is to adjust the sowing Some considerations are necessary, when interpreting 527
472 dates. Adjustment of sowing dates options was explored to results from the scenarios predicted by the GCMs. The most 528

OO
473 investigate a suitable agronomic option for adaptation under significant limitations are their poor resolution, inadequate 529
474 the future climate change scenarios. In this way, for the coupling of atmospheric and oceanic processes, poor simu- 530
475 Cuttack site, the average yield changes of +6.6, +4.1 and lation of cloud processes and inadequate representation of 531
476 9.8% were predicted during July 15 sowing for the GFDL, the biosphere and its feedbacks. The poor resolution is likely 532
477 GISS and UKMO scenarios, respectively, considerably to be significant in northeastern parts of India where the 533
478 higher than those of 19.67, 20.32 and 30.75% observed relief is varied and local climate may be quite different 534

PR
479 during June 15 sowing for the GFDL, GISS and UKMO from the average across the area used by a GCM. Most 535
480 scenarios, respectively. Likewise, the Jorhat site showed GCMs have difficulty in even describing the current climate 536
481 +27.1, +24.3 and +13.4% changes for the sowing on July adequately (Bachelet et al., 1995). The current GCMs are 537
482 1 under the GFDL, GISS and UKMO model scenarios, able to predict neither the changes in the variability of the 538
483 respectively; these changes were considerably higher than weather nor the frequency of catastrophic events such as 539
484 those of +12.13, +12.64 and +8.31% for the June 15 sowing ED hurricanes, floods or even the intensity of monsoons, all of 540
485 for the corresponding scenarios. Further postponement in which can be important in determining crop yields as the 541
486 sowing did not improve the grain yield, probably due to low average climatic data. It seems, therefore, that GCMs can at 542
487 incident solar radiation and temperature. best be used to suggest the likely direction and rate of change 543
488 There were striking differences in the predicted yield of future climates. 544
489 changes among the three scenarios. The large differences According to Long et al. (2005), fertilization effect of 545
CT

490 were mainly due to the sensitivity of spikelet sterility to [CO2] has probably been overestimated, while omission of 546
491 temperature. Even a small difference of just 1 8C could [O3] effects from most models could have led to a 20% 547
492 result in a large yield decrease due to lower number of grains overestimation of crop production in the Northern Hemi- 548
493 being formed (Sheehy et al., 2006). This was illustrated sphere. Database of chamber studies are the mechanistic 549
494 in the two examples used in the present study (Table 6). basis for crop yield models. Hence, these models over- 550
E

495 This was further justified by the current changes in the estimate the yield gain due to elevated [CO2] compared to 551
496 temperature at different sites during the main cropping those observed under fully open-air condition (FACE) 552
RR

497 season (Fig. 2). The imposed climate change scenarios experiments in the field. The current FACE experiments are, 553
498 further enhanced this temperature effect. The increased however, not adequate enough to reparameterize the existing 554
499 spikelet sterility at Cuttack was mainly due to the predicted models (Long et al., 2005). In a recent study, Bannyayan 555
500 increment in temperature above the already high daily et al. (2005) evaluated ORYZA 2000 (Bouman and Van 556
501 maximum temperature (34 8C) (Prasad et al., 2006; Krish- Laar, 2006) against the observed growth and yield of rice in a 557
CO

502 nan and Surya Rao, 2005) under the climate change 3-year field experiment in Japan where rice plants were 558
503 scenarios reaching levels (38 8C) where spikelet damage subjected to the elevated CO2 with FACE under varying N 559
504 was considerable. Although the predicted temperature fertilizer rates. The simulation results showed that the model 560
505 increments were similar to Cuttack, the lower average overestimated the increases in green leaf area indices due to 561
506 temperature (28 8C) at Jorhat was well below the level the elevated CO2 concentration but the enhancement of total 562
507 (30 8C) at which spikelet fertility is affected. At both the biomass was only a minor overestimation. While the model 563
UN

508 locations, the number of spikelets formed was greater under was successful in simulating the increase in rice yield due to 564
509 the changed climates. This could be due to the enhanced the CO2 enrichment, it failed to reproduce the observe 565
510 growth rate of the crop between panicle initiation and interaction with N in the rice yield response to elevated CO2 566
511 flowering as the consequences of fertilizing effect of higher concentration. Thus, the lack of complete understanding 567
512 CO2 level. of the effects and the potential interactions of environment 568
513 The limitation on yields imposed by the increased variables on plant processes preludes the definitive pre- 569
514 spikelet sterility can be largely overcome by the selection of dictions of the effects of global climate change. 570
515 genotypes that possess a higher potential of spikelet fertility Despite the limitations imposed by the assumptions made 571
516 at high temperatures. The fertilizing effect of increased in both the GCM and the crop simulation models, the current 572
517 atmospheric CO2 level is then likely to offset the changes in study provides significant progress in our understanding of 573
518 crop development rate brought about by the increased how future climates are likely to affect rice production in the 574

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019
 AGEE 3025 1–10

10 P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxx–xxx

619
575
574 eastern India. The use of simulation models to predict the Bannyayan, M., Kobayashi, K., Kim, H., Lieffering, M., Okada, M., Mirza, 620
576 likely effects of climate change on crop production is an S., 2005. Modelling the interactive effects of atmospheric CO2 and N on 621
rice growth and yield. Field Crops Res. 93, 237–251. 622
577 evolving process. Our study in view of the findings of the Bouman, B.A.M., Van Laar, H.H., 2006. Description and evaluation of rice 623
578 recent FACE studies clearly shows the need for modification growth model ORYZA 2000 under nitrogen limited conditions. Agric. 624
579 of the existing models. Other levels of production such as the Syst. 87, 249–273. 625
580 influences of water, nutrients and pests, and diseases due to Horie, T., Baskar, J.T., Nakagawa, H., 2000. Crop ecosystem responses to 626
climate change: Rice. In: Reddy, K.R., Hodges, H.F. (Eds.), Climate 627
581 climate change are to be included in the refined models.
change and Global crop productivity. CABI Publishing, Wallingford, 628
582 Some of these limitations in the use of present models can be

F
Oxon, pp. 81–106. 629
583 addressed so that increasingly more accurate predictions can IPCC, 2001. In: Houghton, J.T., Ding, Y., Griggs, D.J., Noguer, M., Van der 630
584 be made in future. Linden, P.J., Da, X., Maskell, K., Johnson, C.A. (Eds.), Climate Change 631

OO
2001, Scientific Basis. Cambridge University Press, New York, USA. 632
Karim, Z., Ahmed, M., Hussain, S.G., Rashid, Kh.B., 1994. Impact of 633
climate change on the production of modern rice in Bangladesh. 634
Uncited references Bangladesh Agricultural Research Council, Dhaka, Bangladesh. 635
585 Kim, H.Y., Lieffering, M., Kobayashi, K., Okada, M., Miura, S., 2003. 636
586 IPCC (2001), Penning de Vries et al. (1989) and Schneider Seasonal changes in the effects of elevated CO2 on rice at three levels of 637
587 (2001). nitrogen supply: A free air CO2 enrichment (FACE) experiment. Global 638

PR
Change Biol. 9, 826–837. 639
Kimbal, B.A., Kobayashi, K., Bindi, M., 2002. Responses of agricultural 640
crops to free air CO2 enrichment. Adv. Agron. 77, 293–368. 641
Acknowledgement Krishnan, P., Surya Rao, A.V., 2005. Effects of genotypic and environmental 642
588 on seed yield and quality of rice. J. Agric. Sci. 143, 283–292. 643
589 Our acknowledgements are to the collaborating members Kropff, M.J., Van Laar, H.H., Mathews, R.B., 1994. ORYZA1: An eco- 644
physiological model for irrigated rice production. In: SARP Research 645
590 from Orissa University of Agriculture & Technology ED Proceedings, IRRI, Wagningen. 646
591 (OUAT), Bhubaneswar; Rice research Station (RRS), Lal, M., Singh, K.K., Rathore, L., Srinivasan, G., Saseendran, S.A., 1998. 647
592 Chinsurah; Narendra Deva University of Agriculture & Vulnerability of rice and wheat yields in NW India to future changes in 648
593 Technology (NDUAT), Faizabad; Jawaharlal Nehru Krishi climate. Agric. Forest Meterol. 89, 101–114. 649
594 Vishwa Vidyalaaya (JNKVV), Jabalpur; Assam Agricultural Long, S.P., Ainsworth, E.A., Leakey, A.D.B., Morgan, P.B., 2005. Global 650
595 University (AAU), Jorhat; Bidan Chandra Krishi Viswa food insecurity. Treatment of major food crops with elevated carbon 651
CT
dioxide or ozone under large-scale fully open-air conditions suggests 652
596 Vidyalaya (BCKV), Kalyani, Rajendra Agricultural Uni- recent models may have over estimated future yields. Phil. Trans. R. 653
597 versity (RAU), Pusa; Indira Gandhi Agricultural University Soc. B 360, 2011–2020. 654
598 (IGAU), Raipur; Birsa Agricultural University (BAU), Mall, R.K., Aggarwal, P.K., 2002. Climate change and rice yields in diverse 655
599 Ranchi; for providing the crop parameter and weather data agro-environments of India. I. Evaluation of impact assessment models. 656
Clim. Change 52, 315–330. 657
600 under the NATP RRPS 25 and Indian Meteorological
E

Mathews, R., Wassmann, R., 2003. Modeling the impacts of climate change 658
601 Department (IMD), Pune, for weather data. and methane emission reductions on rice production: A review. Eur. J. 659
Agron. 19, 573–598. 660
RR

Peng, S.B., Huang, J.L., Sheehy, J.E., Laza, R.C., Visperas, R.M., Zhong, 661
References X., Centeno, G.S., Khush, G.S., Cassman, K.G., 2004. Rice yield 662
602 decline with higher night temperature from global warming. Proc. Natl. 663
Acad. Sci. U.S.A. 101, 9971–9975. 664
603 Aggarwal, P.K., 2003. Impact of climate change on Indian agriculture. J.
604 Plant Biol. 30, 189–198. Penning de Vries, F.W.T., Jansen, D.M., ten Berge, H.F.M., Bakema, A.H., 665
605 Aggarwal, P.K., Kalra, N., Chander, S., Pathak, H., 2004. INFOCROP A 1989. Simulation of ecophysiological processes in several annual crops. 666
CO

606 Generic Simulation model for annual crops in tropical environments. Simulation Monograph. PUDOC, Wagningen, The Netherlands, 271 pp. 667
Penning de Vries, F.W.T., 1993. Rice production and climate change. In: 668
607 IARI, New Delhi, p. 132.
608 Aggarwal, P.K., Kalra, N., Chander, S., Pathak, H., 2006. InfoCrop: A Penning de Vries, F.W.T., Teng, P., Metselaar, K. (Eds.), Systems 669
609 dynamic simulation model for the assessment of crop yields, losses due Approaches for Agricultural Development. Kluwer Academic Publish- 670
ers, Dordrecht, The Netherlands, pp. 175–189. 671
610 to pests, and environmental impact of agro-ecosystems in tropical
611 environments. I. Model description. Agric. Syst. 89 (1), 1–25. Prasad, P.V.V., Boote, K.J., Allen Jr., L.H., Sheehy, J.E., Thomas, J.M.G., 672
2006. Species, ecotype and cultivar differences in spikelet fertility and 673
UN

612 Bachelet, D., Kern, J., Tolg, M., 1995. Balancing the rice carbon budget
613 in China using spatially-distributed data. Ecol. Model 79 (1/3), 167– harvest index of rice in response to high temperature stress. Field Crops 674
Res. 95 (2/3), 398–411. 675
614 177.
615 Baker, J.T., Allen Jr., L.H., 1993. Effects of CO2 and temperature on Schneider, S.H., 2001. What is ‘dangerous’ climate change? Nature 411, 676
616 rice: A summary of five growing seasons. J. Agric. Meterol. 48 (5), 17–19. 677
617 575–582. Sheehy, J.E., Mitchell, P.L., Ferrer, A.B., 2006. Decline in rice grain yields 678
with temperature: Models and correlations can give different estimates. 679
618 Baker, J.T., Allen Jr., L.H., Boote, K.J., 1992. Response of rice to carbon
619 dioxide and temperature. Agric. For. Meterol. 60, 153–166. Field Crops Res. 98, 151–156. 680
680

Please cite this article in press as: Krishnan, P. et al., Impact of elevated CO2 and temperature on rice yield and methods of adaptation as
evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019