Adaptation of crops to climate change through genotypic responses to mean and extreme temperatures

The importance of temperature in the determination of the yield of an annual crop (groundnut; Arachis hypogaea L. in India) was assessed. Simulations from a regional climate model (PRECIS) were used with a crop model (GLAM) to examine crop growth under simulated current (1961-1990) and future (2071-2100) climates. Two processes were examined: the response of crop duration to mean temperature and the response of seed-set to extremes of temperature. The relative importance of, and interaction between, these two processes was examined for a number of genotypic characteristics, which were represented by using different values of crop model parameters derived from experiments. The impact of mean and extreme temperatures varied geographically, and depended upon the simulated genotypic properties. High temperature stress was not a major determinant of simulated yields in the current climate, but affected the mean and variability of yield under climate change in two regions which had contrasting statistics of daily maximum temperature. Changes in mean temperature had a similar impact on mean yield to that of high temperature stress in some locations and its effects were more widespread. Where the optimal temperature for development was exceeded, the resulting increase in duration in some simulations fully mitigated the negative impacts of extreme temperatures when sufficient water was available for the extended growing period. For some simulations the reduction in mean yield between the current and future climates was as large as 70%, indicating the importance of genotypic adaptation to changes in both means and extremes of temperature under climate change. (c) 2006 Elsevier B.V. All rights reserved.

Simulation of the impact of high temperature stress on annual crop yields

Brief periods of high temperature which occur near flowering can severely reduce the yield of annual crops such as wheat and groundnut. A parameterisation of this well-documented effect is presented for groundnut (i.e. peanut; Arachis hypogaeaL.). This parameterisation was combined with an existing crop model, allowing the impact of season-mean temperature, and of brief high-temperature episodes at various times near flowering, to be both independently and jointly examined. The extended crop model was tested with independent data from controlled environment experiments and field experiments. The impact of total crop duration was captured, with simulated duration being within 5% of observations for the range of season-mean temperatures used (20-28 degrees C). In simulations across nine differently timed high temperature events, eight of the absolute differences between observed and simulated yield were less than 10% of the control (no-stress) yield. The parameterisation of high temperature stress also allows the simulation of heat tolerance across different genotypes. Three parameter sets, representing tolerant, moderately sensitive and sensitive genotypes were developed and assessed. The new parameterisation can be used in climate change studies to estimate the impact of heat stress on yield. It can also be used to assess the potential for adaptation of cropping systems to increased temperature threshold exceedance via the choice of genotype characteristics. (c) 2005 Elsevier B.V. All rights reserved.

Design and optimisation of a large-area process-based model for annual crops

The formulation of a new process-based crop model, the general large-area model (GLAM) for annual crops is presented. The model has been designed to operate on spatial scales commensurate with those of global and regional climate models. It aims to simulate the impact of climate on crop yield. Procedures for model parameter determination and optimisation are described, and demonstrated for the prediction of groundnut (i.e. peanut; Arachis hypogaea L.) yields across India for the period 1966-1989. Optimal parameters (e.g. extinction coefficient, transpiration efficiency, rate of change of harvest index) were stable over space and time, provided the estimate of the yield technology trend was based on the full 24-year period. The model has two location-specific parameters, the planting date, and the yield gap parameter. The latter varies spatially and is determined by calibration. The optimal value varies slightly when different input data are used. The model was tested using a historical data set on a 2.5degrees x 2.5degrees grid to simulate yields. Three sites are examined in detail-grid cells from Gujarat in the west, Andhra Pradesh towards the south, and Uttar Pradesh in the north. Agreement between observed and modelled yield was variable, with correlation coefficients of 0.74, 0.42 and 0, respectively. Skill was highest where the climate signal was greatest, and correlations were comparable to or greater than correlations with seasonal mean rainfall. Yields from all 35 cells were aggregated to simulate all-India yield. The correlation coefficient between observed and simulated yields was 0.76, and the root mean square error was 8.4% of the mean yield. The model can be easily extended to any annual crop for the investigation of the impacts of climate variability (or change) on crop yield over large areas. (C) 2004 Elsevier B.V. All rights reserved.

Simulation of crop yields using ERA-40: Limits to skill and nonstationarity in weather-yield relationships

Reanalysis data provide an excellent test bed for impacts prediction systems. because they represent an upper limit on the skill of climate models. Indian groundnut (Arachis hypogaea L.) yields have been simulated using the General Large-Area Model (GLAM) for annual crops and the European Centre for Medium-Range Weather Forecasts (ECMWF) 40-yr reanalysis (ERA-40). The ability of ERA-40 to represent the Indian summer monsoon has been examined. The ability of GLAM. when driven with daily ERA-40 data, to model both observed yields and observed relationships between subseasonal weather and yield has been assessed. Mean yields "were simulated well across much of India. Correlations between observed and modeled yields, where these are significant. are comparable to correlations between observed yields and ERA-40 rainfall. Uncertainties due to the input planting window, crop duration, and weather data have been examined. A reduction in the root-mean-square error of simulated yields was achieved by applying bias correction techniques to the precipitation. The stability of the relationship between weather and yield over time has been examined. Weather-yield correlations vary on decadal time scales. and this has direct implications for the accuracy of yield simulations. Analysis of the skewness of both detrended yields and precipitation suggest that nonclimatic factors are partly responsible for this nonstationarity. Evidence from other studies, including data on cereal and pulse yields, indicates that this result is not particular to groundnut yield. The detection and modeling of nonstationary weather-yield relationships emerges from this study as an important part of the process of understanding and predicting the impacts of climate variability and change on crop yields.

Heat tolerance in groundnut

Tolerance to high soil and air temperature during the reproductive phase is an important component of adaptation to and and semi-arid cropping environments in groundnut. Between 10 and 22 genotypes were screened for tolerance to high air and soil temperature in controlled environments. To assess tolerance to high soil temperature, 10 genotypes were grown from start of podding to harvest at ambient (28 degrees) and high (38 degreesC) soil temperatures, and crop growth rate (CGR), pod growth rate (PGR) and partitioning (ratio PGR:CGR) measured. To assess tolerance to high air temperature during two key stages-microsporogenesis (3-6 days before flowering, DBF) and flowering, fruit-set was measured in two experiments. In the first experiment, 12 genotypes were exposed to short (3-6 days) episodes of high (38 degreesC) day air temperature at 6 DBF and at flowering. In the second experiment, 22 genotypes were exposed to 40 degreesC day air temperature for I day at 6 DBF, 3 DBF or at flowering. Cellular membrane thermostability (relative injury, RI) was also measured in these 22 genotypes. There was considerable variation among genotypes in response to high temperature, whether assessed by growth rates, fruit-set or RI. Pod weight at high soil temperature was associated with variation in CGR rather than partitioning. Flowering was more sensitive to high air temperature than microsporogenesis. Genotypes tolerant to high air temperature at microsporogenesis were not necessarily tolerant at flowering, and nor was tolerance correlated with RI. Six genotypes (796, 55-437, ICG 1236, ICGV 86021, lCGV 87281 and ICGV 92121) were identified as heat tolerant based on their performance in all tests. These experiments have shown that groundnut genotypes can be easily screened for reproductive tolerance to high air and soil temperature and that several sources of heat tolerance are available in groundnut germplasm. (C) 2003 Elsevier Science B.V. All rights reserved.

Drought, pod yield, pre-harvest Aspergillus infection and aflatoxin contamination on peanut in Niger

Soil moisture and soil temperature affect pre-harvest infection with Aspergillus flavus and production of aflatoxin. The objectives of our field research in Niger, West Africa, were to: (i) examine the effects of sowing date and irrigation treatments on pod yield, infection with A. flavus and aflatoxin concentration; and (ii) to quantify relations between infection, aflatoxin concentration and soil moisture stress. Seed of an aflatoxin susceptible peanut cv. JL24 was sown at two to four different sowing dates under four irrigation treatments (rainfed and irrigation at 7, 14 and 21 days intervals) between 1991 and 1994, giving 40 different 'environments'. Average air and soil temperatures of 28-34 degrees C were favourable for aflatoxin contamination. CROPGRO-peanut model was used to simulate the occurrence of moisture stress. The model was able to simulate yields of peanut well over the 40 environments (r(2) = 0.67). In general, early sowing produced greater pod yields, as well as less infection and lower aflatoxin concentration. There were negative linear relations between infection (r(2) = 0.62) and the average simulated fraction of extractable soil water (FESW) between flowering and harvest, and between aflatoxin concentration (r(2) = 0.54) and FESW in the last 25 days of pod-filling. This field study confirms that infection and aflatoxin concentration in peanut can be related to the occurrence of soil moisture stress during pod-filling when soil temperatures are near optimal for A. flavus. These relations could form the basis of a decision-support system to predict the risk of aflatoxin contamination in peanuts in similar environments. (c) 2005 Elsevier B.V. All rights reserved.

Climate change and the flowering time of annual crops

Crop production is inherently sensitive to variability in climate. Temperature is a major determinant of the rate of plant development and, under climate change, warmer temperatures that shorten development stages of determinate crops will most probably reduce the yield of a given variety. Earlier crop flowering and maturity have been observed and documented in recent decades, and these are often associated with warmer (spring) temperatures. However, farm management practices have also changed and the attribution of observed changes in phenology to climate change per se is difficult. Increases in atmospheric [CO2] often advance the time of flowering by a few days, but measurements in FACE (free air CO2 enrichment) field-based experiments suggest that elevated [CO2] has little or no effect on the rate of development other than small advances in development associated with a warmer canopy temperature. The rate of development (inverse of the duration from sowing to flowering) is largely determined by responses to temperature and photoperiod, and the effects of temperature and of photoperiod at optimum and suboptimum temperatures can be quantified and predicted. However, responses to temperature, and more particularly photoperiod, at supraoptimal temperature are not well understood. Analysis of a comprehensive data set of time to tassel initiation in maize (Zea mays) with a wide range of photoperiods above and below the optimum suggests that photoperiod modulates the negative effects of temperature above the optimum. A simulation analysis of the effects of prescribed increases in temperature (0-6 degrees C in + 1 degrees C steps) and temperature variability (0% and + 50%) on days to tassel initiation showed that tassel initiation occurs later, and variability was increased, as the temperature exceeds the optimum in models both with and without photoperiod sensitivity. However, the inclusion of photoperiod sensitivity above the optimum temperature resulted in a higher apparent optimum temperature and less variability in the time of tassel initiation. Given the importance of changes in plant development for crop yield under climate change, the effects of photoperiod and temperature on development rates above the optimum temperature clearly merit further research, and some of the knowledge gaps are identified herein.

Flowering intensity in white yam (Dioscorea rotundata)

White or Guinea yam (Dioscorea rotundata), grown for its underground tubers, is an important food in West Africa. Progress in yam breeding is constrained by variable flowering behaviour, making hybridization difficult. Yam clones may be dioecious, monoecious or hermaphrodite with variable sex ratios. The proportion of plants that flower and the flowering intensity also vary with season and location. The objective of the present work was to investigate whether variation in flowering behaviour was related to factors determining rate of development (photoperiod and temperature through sowing date, location and year) or growth (cumulative solar radiation and temperature). Sex ratios, the proportion of plants that had flower buds and open flowers, and the number of flowers or spikes was recorded in one male (TDr 131) and one female (TDr 99-9) clone of white yam grown in the field in Nigeria at three locations and at different sowing dates. Clone TDr 131 was uniformly male flowering, while clone TDr 99-9 exhibited a number of sex types with gynoecious, monoecious and trimonoecious plants observed. The proportion of flowering plants was low in both clones, averaging 0.34 in clone TDr 131 and 0.13 in clone TDr 99-9. Day of vine emergence had a significant and contrasting effect on the proportion of flowering plants and on flowering intensity in the two clones. In clone TDr 131, the proportion of flowering plants and flowering intensity declined with later vine emergence at all locations (r=0.43-0.53, P<0.05), whereas in clone TDr 99-9 the proportion of flowering plants increased with later emergence (r=0.46, P<0.01). In clone TDr 131, this response was strongly associated with warmer temperatures (r=0.49-0.50; P<0.05) and greater cumulative radiation (r=0.85-0.93; P<0.001) between vine emergence and flowering, rather than photoperiod at vine emergence. This suggests that flowering behaviour in the male clone TDr 131 is strongly influenced by factors that affect growth rather than development. Clone TDr 99-9, on the other hand, exhibited no clear relations between flowering and growth or developmental factors, though the proportion of flowering plants and flowering intensity was greatest at planting dates close to the longest day and at temperatures of 25-26 degrees C. This might suggest that flowering behaviour in clone TDr 99-9 is controlled by photothermal responses.

Duration from vine emergence to flowering suggests a long-day or rate of change of photoperiod response in white yam (Dioscorea rotundata Poir.)

The objective of this study was to quantify the effect of photoperiod on the duration from vine (shoot) emergence to flowering in white or Guinea yam (Dioscorea rotundata). The duration from vine emergence to flowering in two clonal varieties of yam (TDr 131 and TDr 99-9) was recorded at 10 different sowing dates/locations in Nigeria. Durations to flowering varied from 40 to > 88 days. Mean daily temperature and photoperiod between vine emergence and flowering varied from 25 to 27 degrees C and 13.1 to 13.4 h day(-1), respectively. Both clones had similar responses to temperature, with base and optimum temperatures of 12 and 25-27 degrees C, respectively. Thermal durations to flowering were strongly related (r(2) > 0.75-0.83) to absolute photoperiod (h) at vine emergence as well as to rate of change of photoperiod (s day(-1)) at vine emergence. The response to absolute photoperiod suggests that white yams are quantitative LDPs, flowering sooner in long than short days. Yams also flowered earlier when the rate of change of photoperiod was positive but small, or was negative. It is suggested that yams may use a combination of photoperiod and rate of change in order to fine tune flowering time. (c) 2006 Elsevier B.V. All rights reserved.

Phases of dormancy in yam tubers (Dioscorea rotundata)

center dot Background and Aims The control of dormancy in yam (Disocorea spp.) tubers is poorly understood and attempts to shorten the long dormant period (i.e. cause tubers to sprout or germinate much earlier) have been unsuccessful. The aim of this study was to identify and define the phases of dormancy in Dioscorea rotundata tubers, and to produce a framework within which dormancy can be more effectively studied. center dot Methods Plants of 'TDr 131' derived from tissue culture were grown in a glasshouse simulating temperature and photoperiod at Ibadan (7 degrees N), Nigeria to produce tubers. Tubers were sampled on four occasions: 30 d before shoot senescence (149 days after planting, DAP), at shoot senescence (179 DAP), and twice during storage at a constant 25 degrees C (269 and 326 DAP). The development of the apical shoot bud was described from tissue sections. In addition, the responsiveness of shoot apical bud development to plant growth regulators (gibberellic acid, 2-chloroethanol and thiourea) applied to excised tuber sections was also examined 6 and 12 d after treatment. center dot Key Results and Conclusions Three phases of tuber dormancy are proposed: Phase I, from tuber initiation to the appearance of the tuber germinating meristem; Phase II, from the tuber germinating meristem to initiation of foliar primordium; and Phase III, from foliar primordium to appearance of the shoot bud on the surface of the tuber. Phase I is the longest phase (approx. 220 d in 'TDr 131'), is not affected by PGRs and is proposed to be an endo-dormant phase. Phases II and III are shorter (< 70 d in total), are influenced by PGRs and environmental conditions, and are therefore endo-/eco-dormant phases. To manipulate dormancy to allow off-season planting and more than one generation per year requires that the duration of Phase I is shortened.

Inheritance of time to flowering in cowpea (Vigna unguiculata (L.) Walp.)

Time to flowering and maturity is an important adaptive feature in annual crops, including cowpeas (Vigna unguiculata (L.) Walp.). In West and Central Africa, photoperiod is the most important environmental variable affecting time to flowering in cowpea. The inheritance of time from sowing to flowering (f) in cowpeas was studied by crossing a photoperiod-sensitive genotype Kanannnado to a photoperiod-insensitive variety IT97D-941-1. Sufficient seed of F-1, F-2, F-3 and backcross populations were generated. The parental, F-1, F-2, F-3 and the backcross populations were screened for f under long natural days (mean daylength 13.4 h per day) in the field and the parents, F-1, F-2 and backcross populations under short day (10 h per day) conditions. The result of the screening showed that photoperiod in the field was long enough to delay flowering of photoperiod-sensitive genotypes. Photoperiod-sensitivity was found to be partially dominant to insensitivity. Frequency distribution of the trait in the various populations indicated quantitative inheritance. Additive (d) and additive x dominance (j) interactions were the most important gene actions conditioning time to flowering. A narrow sense heritability of 86% was estimated for this trait. This will result in 26 days gain in time to flowering with 5% selection intensity from the F-2 to F-3 generation. At least seven major gene pairs, with an average delay of 6 days each, were estimated to control time to flowering in this cross.

High temperature stress and spikelet fertility in rice (Oryza sativa L.)

In future climates, greater heat tolerance at anthesis will be required in rice. The effect of high temperature at anthesis on spikelet fertility was studied on IR64 (lowland indica) and Azucena (upland Japonica) at 29.6 degrees C (control), 33.7 degrees C, and 36.2 degrees C tissue temperatures. The objectives of the study were to: (i) determine the effect of temperature on flowering pattern; (ii) examine the effect of time of day of spikelet anthesis relative to a high temperature episode on spikelet fertility; and (iii) study the interactions between duration of exposure and temperature on spikelet fertility. Plants were grown at 30/24 degrees C day/night temperature in a greenhouse and transferred to growth cabinets for the temperature treatments. Individual spikelets were marked with paint to relate fertility to the time of exposure to different temperatures and durations. In both genotypes the pattern of flowering was similar, and peak anthesis occurred between 10.30 h and 11.30 h at 29.2 degrees C, and about 45 min earlier at 36.2 degrees C. In IR64, high temperature increased the number of spikelets reaching anthesis, whereas in Azucena numbers were reduced. In both genotypes :511 h exposure to >= 33.7 degrees C at anthesis caused sterility. In IR64, there was no interaction between temperature and duration of exposure, and spikelet fertility was reduced by about 7% per degrees C > 29.6 degrees C. In Azucena there was a significant interaction and spikelet fertility was reduced by 2.4% degrees Cd-1 above a threshold of 33 degrees C. Marking individual spikelets is an effective method to phenotype genotypes and lines for heat tolerance that removes any apparent tolerance due to temporal escape.

Phenotyping parents of mapping populations of rice for heat tolerance during anthesis

Seed set of rice (Oryza sativa L.) is highly sensitive to short episodes of high temperature at anthesis events that are likely to be more frequent in future climates. Breeding for tolerance is therefore an essential component of adaptation to climate variability and change. Experiments were conducted in 2003 and 2004 at optimum (30 degrees C daytime) and high (35 and 38 degrees C) air temperature using parents of some prominent mapping populations (i) to determine whether there were differences in the daily flowering pattern and hence a potential heat avoidance mechanism, and (ii) to identify rice genotypes having true heat tolerance during anthesis, that is, high seed set in spikelets exposed to high temperature. Rice cultivar CG14 (O. glaberrima) reached peak anthesis earlier in the morning (1.5 h after dawn) under both control (30 degrees C) and high (38 degrees C) temperature conditions than O. sativa genotypes (>= 3 h after dawn). Exposure to high temperature (centered on the time of peak anthesis) for 6 h reduced spikelet fertility more than exposure for 2 h, and fertility was lower at 38 degrees C than at 35 degrees C. Genotypic ranking for spikelet fertility at 35 and 38 degrees C was highly correlated in both 2003 and 2004. Fertility was also highly correlated across years, suggesting a consistent and reproducible response of spikelet fertility to temperature. The check cultivar N22 was the most heat tolerant genotype (64-86% fertility at 38 degrees C) and cultivars Azucena and Moroberekan the most susceptible (<8%).