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Key message GWAS on a bread wheat panel with high D genome diversity identified novel alleles and QTLs associated with resilience to combined heat and drought stress under natural field conditions. Abstract As heat (H) and drought stresses occur concurrently under field conditions, studying them separately offers limited opportunities for wheat improvement. Here, a wheat diversity panel containing Aegilops tauschii introgressions was evaluated under H and combined heat–drought (HD) stresses to identify quantitative trait loci (QTLs) associated with resilience to the stresses, and to assess the practicability of harnessing Ae. tauschii diversity for breeding for combined stress resilience. Using genome-wide analysis, we identified alleles and QTLs on chromosomes 3D, 5D, and 7A controlling grain yield (GY), kernel number per spike, and thousand-kernel weight, and on 3D (521–549 Mbp) controlling GY alone. A strong marker–trait association (MTA) for GY stability on chromosome 3D (508.3 Mbp) explained 20.3% of the variation. Leaf traits—canopy temperature, vegetation index, and carbon isotope composition—were controlled by five QTLs on 2D (23–96, 511–554, and 606–614 Mbp), 3D (155–171 Mbp), and 5D (407–413 Mbp); some of them were pleiotropic for GY and yield-related traits. Further analysis revealed candidate genes, including GA20ox, regulating GY stability, and CaaX prenyl protease 2, regulating canopy temperature at the flowering stage, under H and HD stresses. As genome-wide association studies under HD in field conditions are scarce, our results provide genomic landmarks for wheat breeding to improve adaptation to H and HD conditions under climate change.

Heat stress during grain filling has been documented to decrease wheat grain yield and quality in arid regions worldwide. We studied the effect of heat stress on wheat flour quality in heat tolerant cultivars to define the effects of heat stress on flour quality and to identify germplasm combining traits for heat tolerance and good flour quality. We studied the kernel phenotypic traits, the expression of seed storage proteins (SSPs), and the resulting flour quality under heat and normal conditions. Under heat stress, all cultivars yielded narrow-shaped seeds, and increased protein contents as compared to the control plants grown under normal conditions. The specific sedimentation values used to estimate the gluten quality varied between cultivars. We identified cultivars that could maintain good flour quality under heat stress conditions: 'Imam', which possessed the Glu-D1d allele responsible for the suitable bread-making; 'Bohaine', which displayed high expression level of SSPs; and 'Condor', which possessed slight variations in the ratio of each SSP under heat stress conditions. Combining the desirable traits from these cultivars could yield a wheat cultivar with heat tolerance and good flour quality.

Sheep husbandry is considered one of the most important activities in the socio-economic development in the Middle East region, especially in Iraq and Islamic Republic of Iran (IRI). Therefore this study was designed to evaluate the level of ovine pasteurellosis vaccine protective antibody titer and identification of the prevailing serotypes in Iraq (Basrah, Baghdad, Tikrit, Mosul, Erbil). The vaccine was made from pasteurella multosida Bio-type A and the serotypes of Mannheimia haemolytica. This investigation was performed from September 2021 to January 2022, in Iraq. Sheep blood sera samples were obtained from control unvaccinated and vaccinated sheep after 14, 21 and 28 days post vaccination. The results showed that out of 319 sheep blood sera samples which were evaluated using indirect Haemagglutination (IHA) test to detect Mannheimia haemolytica serotypes, the high prevalence (100 %) of M. haemolytica A2 was found in all the five study regions area, while 96.5 % was M. haemolytica A7 and 88.1 % was M. haemolytica A1. The level of antibody titer was measured by specific serum antibody titer of pasteurella multosida Bio-type A. The results revealed that out of 268 vaccinated blood sera samples the overall antibody titer were 12 (3.8 %), 16 (5%) and 17 (5.3 %) for protective antibody titer of 1:160, 1:80 and 1:40 respectively and for antibody titer of 1:20 were 15 (4.7%) and for antibody titer of 1:10 were 17 (5.3 %), whereas the antibody titer in the control group was 4 (7.8 %). The result of this study indicated that the vaccine administered has limited protective power against pasteurella multocida Bio-type A which lead to researchers for further study on identification of specific strain of pasteurella multosida and development of multivalent vaccine including the most prevalent pasteurella serotypes.

Improved nitrogen use-efficient cultivars could be the most economically beneficial and environmentally friendly approach to reduce pollution associated with excessive N fertilization. The performance and genetic gain in grain yield and nitrogen use efficiency (NUE) of a historical set of 12 bread wheat cultivars released for a heat-stressed environment were investigated at four N levels (0 (N0), 43 (N43), 86 (N86), and 129 (N129) kg/ha) for two seasons. Averaged across seasons, increasing N level from N0 to N43, N86, and N129 resulted in yield increases ranging from 4−45%, 13–69%, and 34–87% at N43, N86, and N129, respectively. These yield increases were associated with increases in biomass (r = 0.86, P<0.01). Regressing grain yield of cultivars released during 1960 to 2006 against the year of release showed no trend at N0 and positive nonsignificant trends at N43;. however, significant positive trends were found at N86 and N129 with genetic gain rates of 12.65 and 15.76 kg ha−1 year−1, respectively. This gain was associated with progresses in harvest index (HI) at N43, N86, and N129 but not at N0. On the other hand, during the period from 1960 to 1990, the genetic gain in grain yield at N86 was 24.5 kg ha−1 year−1. Regressing NUE against the year of release showed significant linear trends at N86 and N129 (R2 = 0.511 and R2 = 0.477, respectively), but not at N43. The results indicate that breeders improved grain yield and NUE over 46 years under the heat-stressed environment of Sudan although the rate of increase in yield has been slowed down in recent years. Further improvement in NUE might require broadening the genetic diversity and simultaneous evaluation at low and high N levels.

Wheat (Triticum aestivum L.) is known to be negatively affected by heat stress, and its production is threatened by global warming, particularly in arid regions. Thus, efforts to better understand the molecular responses of wheat to heat stress are required. In the present study, Fourier transform infrared (FTIR) spectroscopy, coupled with chemometrics, was applied to develop a protocol that monitors chemical changes in common wheat under heat stress. Wheat plants at the three-leaf stage were subjected to heat stress at a 42 °C daily maximum temperature for 3 days, and this led to delayed growth in comparison to that of the control. Measurement of FTIR spectra and their principal component analysis showed partially overlapping features between heat-stressed and control leaves. In contrast, supervised machine learning through linear discriminant analysis (LDA) of the spectra demonstrated clear discrimination of heat-stressed leaves from the controls. Analysis of LDA loading suggested that several wavenumbers in the fingerprinting region (400–1800 cm−1) contributed significantly to their discrimination. Novel spectrum-based biomarkers were developed using these discriminative wavenumbers that enabled the successful diagnosis of heat-stressed leaves. Overall, these observations demonstrate the versatility of FTIR-based chemical fingerprints for use in heat-stress profiling in wheat.

PurposeThe purpose of this paper was to study the genetic variability, heritability, heat tolerance indices and phenotypic and genotypic correlation studies for traits of 250 elite International Center for Agricultural Research in the Dry Areas (ICARDA) bread wheat genotypes under high temperature in Wad Medani, Center in Sudan.Design/methodology/approachBread wheat is an important food on a global level and is used in the form of different products. High temperature associated with climate change is considered to be a detrimental stress in the future on world wheat production. A total of 10,250 bread wheat genotypes selected from different advanced yield trials introduction from ICARDA and three checks including were grown in two sowing dates (SODs) (1st and 2nd) 1st SOD heat stress and 2nd SOD non-stress at the Gezira Research Farm, of the Agricultural Research Corporation, Wad Medani, Sudan.FindingsAn alpha lattice design with two replications was used to assess the presence of phenotypic and genotypic variations of different traits, indices for heat stress and heat tolerance for 20 top genotypes and phenotypic and genotypic correlations. Analysis of variance revealed significant differences among genotypes for all the characters. A wide range, 944-4,016 kg/ha in the first SOD and 1,192-5,120 kg/ha in the second SOD, was found in grain yield. The average yield on the first SOD is less than that of the secondnd SOD by 717.7 kg/ha, as the maximum and minimum temperatures were reduced by 3ºC each in the second SOD when compared to the first SOD of the critical stage of crop growth shown.Research limitations/implicationsSimilar wide ranges were found in all morpho-physiological traits studied. High heritability in a broad sense was estimated for days to heading and maturity. Moderate heritability estimates found for grain yield ranged from 44 to 63.6 per cent, biomass ranged from 37.8 to 49.1 per cent and canopy temperature (CT) after heading ranged from 44.2 to 48 per cent for the first and secondnd SODs. The top 20 genotypes are better than the better check in the two sowing dates and seven genotypes (248, 139, 143, 27, 67, 192 and 152) were produced high grain yield under both 1st SOD and 2nd SOD.Practical implicationsThe same genotypes in addition to Imam (check) showed smaller tolerance (TOL) values, indicating that these genotypes had a smaller yield reduction under heat-stressed conditions and that they showed a higher heat stress susceptibility index (SSI). A smaller TOL and a higher SSI are favored. Both phenotypic and genotypic correlations of grain yield were positively and significantly correlated with biomass, harvest index, number of spikes/m2, number of seeds/spike and days to heading and maturity in both SODs and negatively and significantly correlated with canopy temperature before and after heading in both SODs.Originality/valueGenetic variations, heritability, heat tolerance indices and correlation studies for traits of bread wheat genotypes under high temperature

Kernel hardness influences the milling and baking quality of wheat. Stress environments such as heat and combined heat-drought can produce harder kernels, thereby affecting the overall wheat quality. Beside puroindoline genes that are known to determine hardness, other QTLs contribute to the hardness. These QTLs, especially under stress conditions, need extensive research. Moreover, understanding the modification or stabilization of hardness under stress condition and the relationship with stress tolerance will facilitate the selection of superior lines that maintain both high yield and quality even under the stress environment. Therefore, in the current work, we aimed to identify the genetic loci and marker trait associations (MTAs) that contributes for hardness under optimum conditions in Japan, and heat and combined heat-drought (HD) conditions in Sudan. We used a panel of multiple synthetic derivatives (MSD) having diverse Aegilops tauschii genome segments and investigated the association between hardness stabilization and stress tolerance. Under stress conditions, we observed that less reduction of kernel weight is associated with either low change or stable kernel hardness. We identified 47 markers associated with hardness under all conditions, the D genome was the main contributor. For the first time, we found a significant association with hardness under stress conditions on chromosome 4D. We dissected several candidate genes associated with the change of hardness under stress conditions. Our results will improve the understanding of the genetic factors that affect wheat hardness stability.

Aegilops tauschii, the D-genome donor of bread wheat, is a storehouse of genetic diversity that can be used for wheat improvement. This species consists of two main lineages (TauL1 and TauL2) and one minor lineage (TauL3). Its morpho-physiological diversity is large, with adaptations to a wide ecological range. Identification of allelic diversity in Ae. tauschii is of utmost importance for efficient breeding and widening of the genetic base of wheat. This study aimed at identifying markers or genes associated with morpho-physiological traits in Ae. tauschii, and at understanding the difference in genetic diversity between the two main lineages. We performed genome-wide association studies of 11 morpho-physiological traits of 343 Ae. tauschii accessions representing the entire range of habitats using 34,829 DArTseq markers. We observed a wide range of morpho-physiological variation among all accessions. We identified 23 marker&ndash, trait associations (MTAs) in all accessions, 15 specific to TauL1 and eight specific to TauL2, suggesting independent evolution in each lineage. Some of the MTAs could be novel and have not been reported in bread wheat. The markers or genes identified in this study will help reveal the genes controlling the morpho-physiological traits in Ae. tauschii, and thus in bread wheat even if the plant morphology is different.

Twenty-four wheat lines, developed by Aegilops tauschii Coss. introgressions and previously selected for heat or salinity stress tolerance, were evaluated under a drought-rewatering-drought cycle for two years. The objective was to select breeding lines that are resilient to more than one abiotic stress. The experiment was designed in alpha lattice with three replications. Drought was imposed by withholding water during flowering. The results revealed considerable genetic variability in physio-agronomic traits, reflecting the variation in the introgressed segments. High heritability estimates (above 47%) were recorded for most traits, including days to 50% heading, plant height, and thousand-grain weight, indicating the genetic control of these traits which may be useful for cultivar development. The trait-trait correlations within and between water regimes highlighted a strong association among the genetic factors controlling these traits. Some lines exhibited superior performance in terms of stress tolerance index and mean productivity compared with their backcross parent and elite cultivars commonly grown in hot and dry areas. Graphical genotyping revealed unique introgressed segments on chromosomes 4B, 6B, 2D, and 3D in some drought-resilient lines which may be linked to drought resilience. Therefore, we recommend these lines for further breeding to develop climate-resilient wheat varieties.

Heat stress is detrimental to wheat (Triticum aestivum L.) productivity. In this study, we aimed to select heat-tolerant plants from a multiple synthetic derivatives (MSD) population and evaluate their agronomic and physiological traits. We selected six tolerant plants from the population with the background of the cultivar 'Norin 61' (N61) and established six MNH (MSD population of N61 selected as heat stress-tolerant) lines. We grew these lines with N61 in the field and growth chamber. In the field, we used optimum and late sowings to ensure plant exposure to heat. In the growth chamber, in addition to N61, we used the heat-tolerant cultivars 'Gelenson' and 'Bacanora'. We confirmed that MNH2 and MNH5 lines acquired heat tolerance. These lines had higher photosynthesis and stomata conductance and exhibited no reduction in grain yield and biomass under heat stress compared to N61. We noticed that N61 had relatively good adaptability to heat stress. Our results indicate that the MSD population includes the diversity of Aegilops tauschii and is a promising resource to uncover useful quantitative traits derived from this wild species. Selected lines could be useful for heat stress tolerance breeding.

Aegilops tauschii Coss., the D genome donor of hexaploid wheat (Triticum aestivum L.), is the most promising resource used to broaden the genetic diversity of wheat. Taxonomical studies have classified Ae. tauschii into two subspecies, ssp. tauschii and ssp. strangulata. However, molecular analysis revealed three distantly related lineages, TauL1, TauL2 and TauL3. TauL1 and TauL3 includes the only ssp. tauschii, whereas TauL2 includes both subspecies. This study aimed to clarify the phylogeny of Ae. tauschii and to find the traits that can differentiate between TauL1, TauL2 and TauL3, or between ssp. tauschii and ssp. strangulata. We studied the genetic and morpho-physiological diversity in 293 accessions of Ae. tauschii, covering the entire range of the species. A total of 5880 high-quality SNPs derived from DArTseq were used for phylogenetic cluster analyses. As a result, we observed wide morpho-physiological variation in each lineage and subspecies. Despite this variation, no key traits can discriminate lineages or subspecies though some traits were significantly different. Of 124 accessions previously lacking the passport data, 66 were allocated to TauL1, 57 to TauL2, and one to TauL3.

In wheat (Triticum aestivum L.) high temperature (≥30°C) during grain filling leads to considerable reduction in grain yield. We studied 400 multiple synthetic derivatives (MSD) lines to examine the genetic variability of heat stress–adaptive traits and to identify new sources of heat tolerance to be used in wheat breeding programs. The experiment was arranged in an augmented randomized complete block design in four environments in Sudan. A wide range of genetic variability was found in most of the traits in all environments. For all traits examined, we found MSD lines that showed better performance than their parent ‘Norin 61’ and two adapted Sudanese cultivars. Using the heat tolerance efficiency, we identified 13 highly heat-tolerant lines and several lines with intermediate heat tolerance and good yield potential. We also identified lines with alleles that can be used to increase wheat yield potential. Our study revealed that the use of the MSD population is an efficient way to explore the genetic variation in Ae. tauschii for wheat breeding and improvement.

Kernel weight and shape-related traits are inherited stably and increase wheat yield. Narrow genetic diversity limits the progress of wheat breeding. Here, we evaluated kernel weight and shape-related traits and applied genome-wide association analysis to a panel of wheat multiple synthetic derivative (MSD) lines. The MSD lines harbored genomic fragments from Aegilops tauschii. These materials were grown under optimum conditions in Japan, as well as under heat and combined heat–drought conditions in Sudan. We aimed to explore useful QTLs for kernel weight and shape-related traits under stress conditions. These can be useful for enhancing yield under stress conditions. MSD lines possessed remarkable genetic variation for all traits under all conditions, and some lines showed better performance than the background parent Norin 61. We identified 82 marker trait associations (MTAs) under the three conditions, most of them originated from the D genome. All of the favorable alleles originated from Ae. tauschii. For the first time, we identified markers on chromosome 5D associated with a candidate gene encoding a RING-type E3 ubiquitin–protein ligase and expected to have a role in regulating wheat seed size. Our study provides important knowledge for the improvement of wheat yield under optimum and stress conditions. The results emphasize the importance of Ae. tauschii as a gene reservoir for wheat breeding.

Eight chickpea cultivars (Shendi, Jabel Marra, Wad Hamid, Atmor, Hwata, Burgeig, Salwa and Matama) were evaluated for genetic variability, yield stability and contribution of yield attributes to seed yield. Field experiments were carried out for four seasons (2007/2008, 2009/2010, 2010/2011 and 2011/2012) at Hudeiba Research Farm in River Nile State, Sudan. Randomized complete block design with six replications was used. Most of the studied traits recorded highly significant difference (P≤ 0.01) due to cultivars, seasons and their interaction. High heritability and low level of differences among phenotypic coefficient of variation (PCV) and genotypic coefficient of variation (GCV) for studied traits indicated that cultivars influenced more in the expression of these traits. Based on the stability analysis for seed yield; the top yielding cultivars Burgeig and Hwata were adapted to favorable conditions. Both cultivars were late in flowering and maturity and had high number of seeds plant-1, biomass and harvest index. The cultivar Atmor with an intermediate seed yield was the most stable cultivar across seasons. The cultivar Salwa is optional due to its relatively high yield and large seed size. Combining farmer-preferred traits such as high and stable yield, large seed size, plant type and maturity into new cultivars will remain the main objective of the chickpea breeding program in Sudan. Key words: Chickpea cultivars, genetic variability, yield stability, Sudan.

Aims: This experiment aimed at identification of some forage-quality attributes of Sudan pearl millet (Pennisetum glaucum L.) collection for possible future utilization. Study Design: Randomized Complete Block Design (RCBD) with three replications. Place and Duration of Study: The analysis was carried out at the feed analysis laboratory, Elobeid Research Station of Agricultural Research Corporation (ARC), Elobeid, Sudan. Methodology: The selected accessions (42 rainy sown and 38 winter sown) for proximate chemical entities were from 100 pearl millet accessions evaluated for dry matter yield in two field trials. Prior to the proximate quality analysis, the dried samples were ground and oven dried once again. The dried samples were used in 3 replicates for the proximate analysis to determine crude protein (CP), crude fiber (CF) and Nitrogen Free Extract (NFE). Results: Among the studied accessions, nine (in both rainy and winter seasons) had Crude protein (CP%) more than 9%. Nineteen accessions (in rainy) and 36 (in winter) had Crude fiber (CF%) less than 40%, whereas 25 (in rainy) and 32 (in winter) had Nitrogen free extract (NFE%) more than

To accelerate genetic gains in breeding, physiological trait (PT) characterization of candidate parents can help make more strategic crosses, increasing the probability of accumulating favorable alleles compared to crossing relatively uncharacterized lines. In this study, crosses were designed to complement “source” with “sink” traits, where at least one parent was selected for favorable expression of biomass and/or radiation use efficiency—source—and the other for sink-related traits like harvest-index, kernel weight and grains per spike. Female parents were selected from among genetic resources—including landraces and products of wide-crossing (i.e. synthetic wheat)—that had been evaluated in Mexico at high yield potential or under heat stress, while elite lines were used as males. Progeny of crosses were advanced to the F4 generation within Mexico, and F4-derived F5 and F6 generations were yield tested to populate four international nurseries, targeted to high yield environments (2nd and 3rd WYCYT) for yield potential, and heat stressed environments (2nd and 4th SATYN) for climate resilience, respectively. Each nursery was grown as multi-location yield trials. Genetic gains were achieved in both temperate and hot environments, with most new PT-derived lines expressing superior yield and biomass compared to local checks at almost all international sites. Furthermore, the tendency across all four nurseries indicated either the superiority of the best new PT lines compared with the CIMMYT elite checks, or the superiority of all new PT lines as a group compared with all checks, and in some cases, both. Results support—in a realistic breeding context—the hypothesis that yield and radiation use efficiency can be increased by improving source:sink balance, and validate the feasibility of incorporating exotic germplasm into mainstream breeding efforts to accelerate genetic gains for yield potential and climate resilience.

Adaptation of wheat (Triticum aestivum L.) to high temperatures could be improved by introducing alien genes from wild relatives. We evaluated the responses of wheat-Leymus racemosus chromosome introgression lines to high temperature to determine their potentiality for developing improved wheat cultivars. Introgression lines and their parent Chinese Spring were evaluated in a growth chamber at the seedling stage and in the field at the reproductive stage in two heat-stressed environments in Sudan. Optimum and late planting were used to ensure exposure of the plants to heat stress at the reproductive stage. The results revealed the impact of several Leymus chromosomes in improving wheat adaptation and tolerance to heat. Three lines possessed enhanced adaptation, whereas two showed high heat tolerance. Two addition lines showed a large number of kernels per spike, while one possessed high yield potential. Grain yield was correlated negatively with the heat susceptibility index, days to heading and maturity and positively with kernel number per spike and triphenyl tetrazolium chloride assay under late planting. The findings suggest that these genetic stocks could be used as a bridge to introduce the valuable Leymus traits into a superior wheat genetic background, thus helping maximize wheat yield in heat-stressed environments.

Understanding wheat (Triticum aestivum L.) response to high shoot/root temperature during the early growth stages is important for successful production in tropical and subtropical environments. This study examined the physiological response of wheat cultivars to high shoot and/or root temperatures during early growth stages. Three cultivars; Imam, Fang and Siete Cerros were grown in soil and hydroponically at three shoot/root temperatures (23/23, 23/35 and 35/35ºC for the soil experiment; and 22/22, 22/38 and 38/38ºC for the hydroponic experiment). Leaf dry weight and leaf area plant-1 were significantly decreased by high shoot/ root temperature (HS/HR, 35/35 and 38/38ºC) but was not affected by a normal shoot/high root temperature (NS/HR, 23/35 and 22/38ºC). The NS/HR (22/38ºC) and HS/HR (38/38ºC) treatments in the hydroponic experiment significantly decreased photosystem II quantum yield ( Φpsii), photosynthetic rate (Pn) and specific leaf area (SLA) compared with the normal shoot/normal root (NS/NR, 22/22ºC) temperature treatment. Chlorophyll accumulation was significantly decreased by NS/HR, but increased significantly by HS/HR in most of the measuring dates. The heat-tolerant cultivar, Fang, always had the highest chlorophyll content, Φpsii and Pn under all temperature treatments, while the heat-sensitive cultivar, Siete Cerros, always had the greatest reduction in these traits especially towards the end of the experiment. Imam and Fang responded to HS/HR in the hydroponic experiment by immediate and greater reductions in leaf dry weight, total leaf area and SLA during the first wk of the treatments compared with Siete Cerros. The response changed with the treatments duration such that Imam showed the least reduction and Siete Cerros was the most affected cultivar towards the end of the experiment. Thus, wheat cultivars differentially responded to high shoot/root temperature by reducing the leaf weight and area and hence accumulating more chlorophyll in the diminished leaves. The failure to undergo such changes led to significantly lower chlorophyll accumulation, Φpsii and Pn under high root temperature.

High temperatures limit wheat (Triticum aestivum L.) production in many areas around the world. Soil temperatures near the root zone could be as high as the air temperature during wheat grain filling. The objectives of this study were to investigate the effects of high soil temperature on grain yield and related traits of wheat genotypes and to examine their differential responses. Three genotypes, Imam, Fang and Siete Cerros were grown under three temperature conditions in the daytime during grain-filling period: (i) normal air temperature/normal soil temperature (26/26°C), (ii) normal air temperature/high soil temperature (26/38°C) and (iii) high air temperature/high soil temperature (38/38°C). The night temperature was 18/18°C in all treatments. Temperatures during the daytime were increased at a rate of 4°C hr-1 from 18°C to the designated temperature, which was kept for at least 4 hr around noon. The 26/38°C and 38/38°C treatments significantly decreased the chlorophyll content (SPAD) of flag leaves, grain-filling duration, and increased carbohydrate remobilization or loss from the stem and the root, but with varying degrees among genotypes. Grain yield, biomass, grain weight, grains number spike-1 and harvest index at the 38/38°C treatment were significantly lower than at the other two treatments. In Imam, the grain yield was lower at 26/38°C than at 26/26°C, while in Siete Cerros the grain yield, grain weight, grain number spike-1 and harvest index were lower at the 38/38°C treatment than at the other two treatments. These results indicated that high temperature of soil alone (26/38°C) or high temperature of both air and soil (38/38°C) decreased the chlorophyll content and grain-filling duration, and increased carbohydrate remobilization. Genotypic differences in the responses to high soil temperature (26/38°C) and high air/soil temperature (38/38°C) were also observed.