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    Impact of elevated CO2 and temperature on rice yield and methodsof adaptation as evaluated by crop simulation studies

    P. Krishnan a,*, D.K. Swain b, S. Chandrabaskar b, S.K. Nayaka, R.N. Dash b

    aDivision of Biochemistry, Plant Physiology and Environmental Sciences, Central Rice Research Institute, Cuttack 753006, India

    bDivision 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

    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 of7.20 and 6.66%, respectively, atthe 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 of9.02, 11.30 and 21.35%. There were considerabledifferences 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 Published by Elsevier B.V.

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

    1. Introduction

    The climatic variability and the predictedclimatic changes

    are of major concern to the rice crop scientists because of

    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 incre-

    asing 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 theabsenceof temperature increase, many studies have shown that the net

    effect of doubling of CO2 was increase in the yield of rice

    (Kimetal.,2003). It becomes necessaryto assessthe 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.,

    www.elsevier.com/locate/ageeAgriculture, Ecosystems and Environment xxx (2007) xxxxxx

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    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: [email protected] (P. Krishnan).

    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

    mailto:[email protected]://dx.doi.org/10.1016/j.agee.2007.01.019http://dx.doi.org/10.1016/j.agee.2007.01.019http://dx.doi.org/10.1016/j.agee.2007.01.019http://dx.doi.org/10.1016/j.agee.2007.01.019mailto:[email protected]
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    1992; 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 increasewas 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 pheno-

    logical development are calculated. ORYZA1 calculates dry

    matter production as a function of light, CO2 and tempe-

    rature 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 calcula-

    tions of the distribution of light within the canopies,

    the radiation absorbed by the canopy and photosynthesis

    light response cure of leaves. Growth and maintenance

    respiration are calculated as a function of tissue N content,

    temperature and crop-specific coefficients. This methodol-

    ogy, although yields very accurate results, poses practical

    difficulties because of its requirement for detailed and

    careful measurements (Kropff et al., 1994). The INFOCROPmodel, 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 develop-

    ment stage (Aggarwal et al., 2004).

    The rice-growing regions included for the present study

    lie in eastern India, and these sites (Bhubaneswar, Chin-

    surah, 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 20012004 (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 xxx (2007) xxxxxx2

    AGEE 3025 110

    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

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

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    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 inverse-

    distance-squared algorithm to a 0.58 latitude 0.58 long-itude grid using a raster-based Geographical Information

    Systems (GIS) software package. Scenarios were produced

    by applying ratios of precipitation or differences in tempe-

    rature predicted for the 2 CO2 and 1 CO2 simulation tothe 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 forthe 1 CO2 (380 ppm) and 2 CO2 (760 ppm) simulationsto the baseline daily weather data set. The changes between

    1 CO2 and 2 CO2 conditions were representative of thedifferences 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

    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, 20022003). 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

    of7.20% in yields for every 1 8C increase in temperature,while INFOCROP predicted 6.66%. But increasing CO2concentration (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, 9.38and 15.86% in yield for the GDFL, GISS and UKMOscenarios, respectively. For the corresponding scenarios,

    INFOCROP indicated larger reductions at 9.02, 11.30and 21.35%, respectively. When each site was analysedindividually 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,

    P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxxxxx4

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    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

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    Table 2c

    Model inputs of geographical information of different locations

    Soil properties Bhubaneswar Kalyani Faizabad Ranchi Pusa Raipur Jabalpur Jorhat Cuttack

    Latitude 208140N 228570N 228870N 238170N 22850N 258590N 238090N 26880N 208300N

    Longitude 858520E 888210E 88840E 858190E 868E 85840E 798580E 95850 E 868E

    Altitude (m) 25.9 7.8 8.62 625 23 51.84 411 86.56 23

    Table 3

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

    GFDL GISS UKMO

    Source laboratory Geophysical Fluid Dynamics

    Laboratory

    Goddard Institute for

    Space Studies

    United Kingdom

    Meteorological Office

    Base 1 CO2 (ppm) 380 380 380Change in global temperature (8C) +4.0 +4.2 +5.2

    Change in global precipitation (%) 8 11 15

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

    AGEE 3025 110

    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

    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 20.39400 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 4.85600 40.61 29.39 16.74 1.04 15.80 29.17 7.14700 56.37 41.21 27.96 9.85 7.67 23.15 17.43Average 25.14 16.86 3.65 10.56 23.59 34.05 3.76

    ORYZA

    380 0 4.27 15.32 26.78 29.45 38.96 19.13400 2.78 0.93 8.44 19.74 23.67 33.65 13.22500 18.57 11.73 0.24 11.75 15.43 26.57 3.87600 26.8 18.54 5.03 6.86 10.87 21.84 1.80700 30.73 23.77 8.67 3.68 7.32 19.04 5.52Average 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

    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

    (t/ha)

    GFDL GISS UKMO

    Predicted

    change (%)

    Predicted

    yield (t/ha)

    Predicted

    change (%)

    Predicted

    yield (t/ha)

    Predicted

    change (%)

    Predicted

    yield (t/ha)

    Bhubaneswar 4.46 23.87 3.40 27.45 3.24 37.22 2.80Chinsurah 5.18 7.03 4.82 7.38 4.80 8.11 4.76Cuttack 4.93 25.44 3.68 27.67 3.57 40.87 2.92Faizabad 4.72 13.55 4.08 17.65 3.89 28.34 3.38Jabalpur 7.54 10.7 6.73 14.04 6.48 25.66 5.61Jorhat 3.83 13.51 4.35 12.32 4.30 7.55 4.12

    Kalyani 3.55 8.73 3.24 11.65 3.14 22.38 2.76Pusa 3.82 3.74 3.68 4.35 3.65 5.26 3.62Raipur 3.75 1.71 3.69 5.11 3.56 18.01 3.07Ranchi 4.5 8.89 4.10 12.01 3.96 35.15 2.92Average 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.

    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

    (t/ha)

    GFDL GISS UKMO

    Predictedchange (%)

    Predictedyield (t/ha)

    Predictedchange (%)

    Predictedyield (t/ha)

    Predictedchange (%)

    Predictedyield (t/ha)

    Bhubaneswar 4.46 17.33 3.69 20.36 3.55 27.53 3.23Chinsurah 5.18 8.03 4.76 8.72 4.73 9.59 4.68Cuttack 4.93 19.67 3.96 20.32 3.93 30.75 3.41Faizabad 4.72 9.02 4.29 11.27 4.19 18.82 3.83Jabalpur 7.54 11.05 6.71 14.08 6.48 21.05 5.95Jorhat 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 2.96Pusa 3.82 4.93 3.63 6.31 3.58 6.58 3.57Raipur 3.75 2.79 3.65 5.22 3.55 10.09 3.37Ranchi 4.50 7.87 4.15 10.35 4.03 25.98 3.33Average 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.

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    to describe the response of spikelet fertility to temperature

    is

    d 100

    1 e0:853TmaxTmp

    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, Tmphad 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 availableweather 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

    tolerance of the variety by not adjusting the value of Tmp,

    large decreases in yield due to spikelet sterility were pre-

    dicted. 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 andUKMO 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. Theseresults 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., 1990

    a,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 assimila-

    tion rate and canopy net photosynthesis also increased with

    increasing CO2 concentration. The elevated CO2 concentra-

    tion was found to accelerate the development but shorten the

    total growth duration of rice.

    There are many indirect influences of elevated CO2on 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 Lincon, 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 taken

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

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    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

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    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.

    Fig. 4. Changes in yield (%) under the GCMs scenarios of the rice variety

    IR 36 under different sowing dates grown during the kharifseason at Jorhat.

    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.

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    these factors into account, there are still limitations on their

    predictive value.

    The average yield changes of 8.23 and 7.31% byORYZA and INFOCROP, respectively, due to the effect

    of temperature when simulated on per degree Celsius

    basis, were comparable with that of10% 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%) clearlyshow 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 max-

    imum 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 concen-

    tration. 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). Themain 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 charac-

    terized 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 phenologicaldevelopment 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 overallregional rice yield. Averaged across all three GCM scena-

    rios, 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 tempera-

    tures.Among the different sites tested, both the models pre-

    dicted 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 sub-

    humid 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 a n d a

    minimum temperature of 19 8C only, which probably contri-

    buted to the benefits from the predicted effects of climate

    change scenarios. The predicted declines in the overall rice

    P. Krishnan et al. / Agriculture, Ecosystems and Environment xxx (2007) xxxxxx8

    AGEE 3025 110

    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

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    yield by both cop models for the GFDL, GISS 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 offsetany 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 of19.67,20.32 and30.75% observedduring 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; Krish-

    nan and Surya Rao, 2005) under the climate change

    scenarios reaching levels ($38 8C) where spikelet damagewas 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

    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 riceproduction.

    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 simu-

    lation 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 likelydirection 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 Hemi-

    sphere. Database of chamber studies are the mechanistic

    basis for crop yield models. Hence, these models over-estimate 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 CO2concentration. Thus, the lack of complete understanding

    of the effects and the potential interactions of environment

    variables on plant processes preludes the definitive pre-

    dictions 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

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

    AGEE 3025 110

    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

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    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

    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.

    Uncited references

    IPCC (2001), Penning de Vries et al. (1989) and Schneider

    (2001).

    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 Uni-

    versity (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 MeteorologicalDepartment (IMD), Pune, for weather data.

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    evaluated by crop simulation studies, Agric. Ecosyst. Environ. (2007), doi:10.1016/j.agee.2007.01.019

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