Whole plant responses, key processes, and adaptation to drought stress: the case of rice

Most high-yielding rice cultivars developed for irrigated conditions, including the widely grown lowland variety IR64, are highly susceptible to drought stress. This limits their adoption in rainfed rice environments where there is a risk of water shortage during the growing season. Mapping studies using lowland-by-upland rice populations have provided limited information about the genetic basis of variation in yield under drought. One approach to simultaneously improve and understand rice drought tolerance is to generate backcross populations, select superior lines in managed stress environments, and then evaluate which features of the selected lines differ from the recurrent parent. This approach was been taken with IR64, using a range of tolerant and susceptible cultivars as donor parents. Yields of the selected lines measured across 13 widely contracting water environments were generally greater than IR64, but genotype-by-environment effects were large. Traits expected to vary between IR64 and selected lines are plant height, because many donors were not semi-dwarf types, and maturity, because selection in a terminal stress environment is expected to favour earliness. In these experiments it was found that some lines that performed better under upland drought were indeed taller than IR64, but that shorter lines with good yield under drought could also be identified. In trials where drought stress developed in previously flooded (lowland) fields, height was not associated with performance. There was little change in maturity with selection. Other notable differences between IR64 and the selected backcross lines were in their responses to applied ABA and ethylene in greenhouse experiments at the vegetative stage and in leaf rolling observed under chronic upland stress in the field. These observations are consistent with the hypothesis that adaptive responses to drought can effectively allow for improved performance across a broad range of water environments. The results indicate that the yield of IR64 under drought can be significantly improved by backcrossing with selection under stress. In target environments where drought is infrequent but significant in certain years, improved IR64 with greater drought tolerance would be a valuable option for farmers.     

Expression of the CCCH‐tandem zinc finger protein gene OsTZF5 under a stress‐inducible promoter mitigates the effect of drought stress on rice grain yield under field conditions

Increasing drought resistance without sacrificing grain yield remains an ongoing challenge in crop improvement. In this study, we report that O ryza s ativa CCCH‐t andem z inc f inger protein 5 (OsTZF5) can confer drought resistance and increase grain yield in transgenic rice plants. Expression of OsTZF5 was induced by abscisic acid, dehydration and cold stress. Upon stress, OsTZF5‐GFP localized to the cytoplasm and cytoplasmic foci. Transgenic rice plants overexpressing OsTZF5 under the constitutive maize ubiquitin promoter exhibited improved survival under drought but also growth retardation. By introducing OsTZF5 behind the stress‐responsive OsNAC6 promoter in two commercial upland cultivars, Curinga and NERICA4, we obtained transgenic plants that showed no growth retardation. Moreover, these plants exhibited significantly increased grain yield compared to non‐transgenic cultivars in different confined field drought environments. Physiological analysis indicated that OsTZF5 promoted both drought tolerance and drought avoidance. Collectively, our results provide strong evidence that OsTZF5 is a useful biotechnological tool to minimize yield losses in rice grown under drought conditions.     

Markers associated with a QTL for grain yield in wheat under drought

Drought is a major abiotic stress that adversely affects wheat production in many regions of the world. The objective of this study was to identify quantitative trait loci (QTL) controlling grain yield and yield components under reduced moisture. A cross between common wheat cultivars ‘Dharwar Dry’ (drought tolerant) and ‘Sitta’ was the source of one hundred twenty-seven recombinant inbred lines evaluated for two-seasons in a field under differing soil moisture regimes in Ciudad Obregon, Sonora, Mexico. An SSR/EST-STS marker map was constructed and a grain yield QTL on the proximal region of chromosome 4AL was found to have a significant impact on performance under reduced moisture. This region was associated with QTL for grain yield, grain fill rate, spike density, grains m⁻², biomass production, biomass production rate, and drought susceptibility index (DSI). Molecular markers associated with these traits explained 20, 33, 15, 23, 30, 26, and 41% of phenotypic variation, respectively on chromosome 4A. Microsatellite locus Xwmc89 was associated with all significant QTL covering a 7.7 centiMorgans (cM) region and generally explained the greatest proportion of phenotypic variation. The alleles associated with enhanced performance under drought stress were contributed by Dharwar Dry. Microsatellite marker wmc89 may be useful for marker assisted selection to enhance drought tolerance.     

Assessment of grain yield indices in response to drought stress in wheat (Triticum aestivum L.)

Drought stress constricts crop production in the world. Increasing human population and predicted temperature increase owing to global warming will lead ruthless problems for agricultural production in near future. Hence, use of high yielding genotypes having drought tolerance and scrutinize of drought sensitive local cultivars for making them tolerant may be the proficient approaches to cope its detrimental outcomes. The current study was executed during 20015–2016 and 2016–2017 in field using randomized complete block design under factorial arrangements on 50 wheat genotypes for exploring their sensitivity and tolerance against drought. Some of the attributes of grain yield and drought tolerance indices were recorded. Grain yield showed negative correlations with tolerance index (TOL), drought index (DI) and stress susceptibility index (SSI) while positive correlation with mean productivity (MP) and geometric mean productivity (GMP) under drought condition. These findings depicted that tolerant genotypes could be chosen by high MP and GMP values and low SSI and TOL values. Based on the results, genotypes GA-02, Faisalabad-83, 9444, Sehar-06, Pirsabak-04 and Kohistan-97 were more tolerant and recognized as suitable for both normal and drought conditions. Genotypes of Chenab-00, Kohsar-95, Parwaz-94 and Kohenoor-83 confirmed more sensitive due to high grain yield loss under drought stress.     

Identification of quantitative trait loci for four morphologic traits under water stress in rice （Oryza sativa L.）

Late season drought coinciding with the rice booting to heading stage affects the development of plant height,panicle exsertion,and flag leaf size,and causes significant yield loss.In this study,a recombinant inbred line population derived from a cross between paddy and upland cultivars was used for data collection of the morphologic traits under well water and drought stress conditions.bought stress was applied at the stage of panicle initiation in the field in 2002 and at the booting stage in PVC pipes in 2003.The data from stress con ditions and their ratios(tait measured under stress condition/trait measured under well water condition)or differences(trait measured under stress condition minus trait measured under well water condition)were used for OTL analysis.Totally,17 and 36 QTLs for these traits were identified in 2002 and 2003,respectively,which explained a range of 2.58%-29.82%Of the phenotypic variation.Among them,six QTLs were commonly identified in the two years,suggesting that the drought stress in the two years was different.The genetic basis of these traits will provide useful information for improving rice late season drought resistance,and their application as indirect indices in rice late season drought resistance screening was also discussed.     

Expression profiling of rice cultivars differing in their tolerance to long-term drought stress

Understanding the molecular basis of plant performance under water-limiting conditions will help to breed crop plants with a lower water demand. We investigated the physiological and gene expression response of drought-tolerant (IR57311 and LC-93-4) and drought-sensitive (Nipponbare and Taipei 309) rice (Oryza sativa L.) cultivars to 18 days of drought stress in climate chamber experiments. Drought stressed plants grew significantly slower than the controls. Gene expression profiles were measured in leaf samples with the 20 K NSF oligonucleotide microarray. A linear model was fitted to the data to identify genes that were significantly regulated under drought stress. In all drought stressed cultivars, 245 genes were significantly repressed and 413 genes induced. Genes differing in their expression pattern under drought stress between tolerant and sensitive cultivars were identified by the genotype x environment (G x E) interaction term. More genes were significantly drought regulated in the sensitive than in the tolerant cultivars. Localizing all expressed genes on the rice genome map, we checked which genes with a significant G x E interaction co-localized with published quantitative trait loci regions for drought tolerance. These genes are more likely to be important for drought tolerance in an agricultural environment. To identify the metabolic processes with a significant G x E effect, we adapted the analysis software MapMan for rice. We found a drought stress induced shift toward senescence related degradation processes that was more pronounced in the sensitive than in the tolerant cultivars. In spite of higher growth rates and water use, more photosynthesis related genes were down-regulated in the tolerant than in the sensitive cultivars.     

Identification and characterization of large-effect quantitative trait loci for grain yield under lowland drought stress in rice using bulk-segregant analysis

An F_4:5 population of 490 recombinant inbred lines (RILs) from the cross Apo/^2*Swarna was used to detect quantitative trait loci (QTL) with large effects on grain yield under drought stress using bulk-segregant analysis (BSA). Swarna is an important rainfed lowland rice variety grown on millions of hectares in Asia, but is highly susceptible to drought and aerobic soil conditions. Apo is an aerobic-adapted variety with moderate tolerance to drought. Two rice microsatellite (RM) markers, RM324, and RM416, located on chromosomes 2 and 3, respectively, were shown via BSA to be strongly associated with yield under lowland drought stress. The effects of these QTL were tested in a total of eight hydrological environments over a period of 3 years. The QTL linked to RM416 (DTY _3.1 ) had a large effect on grain yield under severe lowland drought stress, explaining about 31% of genetic variance for the trait (P < 0.0001). It also explained considerable variance for yield under mild stress in lowland conditions and aerobic environments. To our knowledge this is the first reported QTL that has a large effect on yield in both lowland drought and aerobic environments. The QTL linked to RM324 (DTY _2.1 ) had a highly significant effect on grain yield in lowland drought stress (R ² = 13–16%) and in two aerobic trials. The effect of these QTL on grain yield was verified to be not mainly due to phenology differences. Effects of DTY _3.1 on yield under stress have been observed in several other rice mapping populations studied at IRRI. Results of this study indicate that BSA is an effective method of identifying QTL alleles with large effects on rice yield under severe drought stress. The Apo alleles for these large-effect QTL for grain yield under drought and aerobic conditions may be immediately exploited in marker-assisted-breeding to improve the drought tolerance of Swarna.     

Cereal flag leaf adaptations for grain yield under drought: knowledge status and gaps

Selection for drought-tolerant cereal varieties has successfully moved to screening for grain yield under stress. Grain yield is the culmination of the process of grain filling, which in turn is closely linked to flag leaf functionalities. For grain filling to occur under drought, either a relatively uncompromised or a favorably reprogrammed functioning of the flag leaf is required. However, knowledge is limited on how effectively flag leaves can function under stress conditions or what adaptations could allow such functioning. The information on rice flag leaf function and/or adaptation under drought is critically limited, while rice continues to be the crop with the highest potential to alleviate hunger and poverty. In fact, other cereal crops are equally important in maintaining regional food baskets and these too suffer intermittently from different intensities and kinds of drought. Patchy information is available on the morpho-anatomical, physiological and biochemical aspects of flag leaves under drought; even this is dispersed within different cereals, with studies predominantly on wheat. Hence, a reasonable understanding of the function of flag leaf under drought is lacking for any cereal. Importantly, very few reports exist on the molecular and mechanistic understanding of any known adaptations of flag leaf function under drought. Here we review the existing information on cereal flag leaf function under drought and highlight the need to better understand its characteristics/adaptations, especially at the molecular level. Novel drought-tolerant breeding material generated through selection for yield under stress can be a useful resource to underpin the mechanistic basis of the contribution of flag leaves to such yield. Improved knowledge can then be used for providing dependable markers (morphological, anatomical, physiological, biochemical and/or molecular) for robust flag leaves, leading to efficient and judicious use of resources for screening broader germplasm collections.     

Soybean cultivar variability for nodule formation and growth under drought

Both nodulation and nitrogen fixation in soybean [Glycine max (L.) Merr.] are sensitive to soil drying, which can have important negative effects on yield. An exception to this general response has been the identification of the cultivar Jackson as being drought tolerant for N₂ fixation. The objectives of this research were to examine nodule formation and growth in Jackson among other soybean cultivars in response to soil drying under field conditions. Two field experiments were conducted to examine the genetic variation in the sensitivity of nodule numbers and dry weights to soil drying. Substantial variation among soybean lines was found, and the drought-tolerance trait was demonstrated again in Jackson. Greenhouse experiments were conducted to further analyze the variation of nodulation response to soil water content. The differences among cultivars observed in the field were confirmed in the greenhouse. Importantly, the relative drought insensitivity of N₂ fixation in cultivar Jackson was associated with high individual nodule dry weight under drought conditions, relative to well-watered plants. It was concluded that large variation in nodulation sensitivity to water deficit exists among soybean cultivars and that the response of N₂ fixation rates to drought is related in part to nodule formation and growth.     

Evaluating the resistance of six rice cultivars to drought: restriction of deep rooting and the use of raised beds

Soil water deficits reduce rice (Oryza sativa L.) productivity under upland field conditions. In this study, we constructed screening facilities to evaluate the performance of rice cultivars under drought conditions and to assess the roles of deep roots. Two experiments were conducted with six rice cultivars, including drought-tolerant and drought-susceptible cultivars, grown in two root environments: a root-restricted treatment that restricted rooting depth with water-permeable sheets, and a raised bed that reduced water availability in the surface soil by inserting a gravel layer between the topsoil and subsoil layers to interrupt capillary transport of water. In the root-restricted treatment, in which root growth was restricted to the surface 25-cm layer, leaf water potential decreased faster in cultivars with a large canopy during drought stress, and there was little difference in panicle weight among cultivars. With a normal (unrestricted) root environment, the deepest-rooting cultivar (‘IRAT109’) maintained higher leaf water potential during drought, although panicle weight under drought stress was affected by yield potential as well as by deep rooting. Under the intermittent drought stress in the raised bed, deep-rooting cultivars accumulated more nitrogen and produced more biomass, and the difference in panicle weight between deep-rooting drought-tolerant and shallow-rooting drought-susceptible cultivars was magnified by the raised bed compared with the yield differences under drought in a normal root environment. These results demonstrate that the drought screening facilities we developed can help to identify superior cultivars under upland field conditions without time-consuming measurement of deep root systems.     

A large-effect QTL for rice grain yield under upland drought stress on chromosome 1

Drought is a major abiotic stress factor limiting rice production in rainfed areas. In this study we identified a large-effect quantitative trait locus (QTL) associated with grain yield under stress in five different populations on chromosome 1. The effect of this QTL was further confirmed and characterized in five backcross populations in a total of sixteen stress and non-stress trials during 2006 and 2008. In all the stress trials (eight in total) qDTY1.1 showed strong association with grain yield explaining on average 58% of the genetic variation in the trait. Homozygotes for the tolerant parent allele (Apo) yielded on average 27% more than the susceptible parent allele (IR64) homozygotes. Using an Apo/^3*IR64 population, the peak of this QTL (qDTY1.1) was mapped to an interval between RM486 and RM472 at 162.8?cM at a LOD score of 9.26. qDTY1.1 was strongly associated with plant height in all the environments; this was probably due to the presence of the sd1 locus in this genomic region. In a Vandana/^3*IR64 population segregating for sd1, a strong relation between plant height and yield under stress was observed. The observed relation between increased height and drought tolerance is likely due to tight linkage between qDTY1.1 and sd1 and not due to pleiotrophy of sd1. Thus there is a possibility of combining reduced plant height and drought tolerance in rice. The large and consistent effect of qDTY1.1 across several genetic backgrounds and environments makes it a potential strong candidate for use in molecular breeding of rice for drought tolerance.     

Overexpression of an Arabidopsis thaliana galactinol synthase gene improves drought tolerance in transgenic rice and increased grain yield in the field

Drought stress has often caused significant decreases in crop production which could be associated with global warming. Enhancing drought tolerance without a grain yield penalty has been a great challenge in crop improvement. Here, we report the Arabidopsis thaliana galactinol synthase 2 gene (AtGolS2) was able to confer drought tolerance and increase grain yield in two different rice (Oryza sativa) genotypes under dry field conditions. The developed transgenic lines expressing AtGolS2 under the control of the constitutive maize ubiquitin promoter (Ubi:AtGolS2) also had higher levels of galactinol than the non‐transgenic control. The increased grain yield of the transgenic rice under drought conditions was related to a higher number of panicles, grain fertility and biomass. Extensive confined field trials using Ubi:AtGolS2 transgenic lines in Curinga, tropical japonica and NERICA4, interspecific hybrid across two different seasons and environments revealed the verified lines have the proven field drought tolerance of the Ubi:AtGolS2 transgenic rice. The amended drought tolerance was associated with higher relative water content of leaves, higher photosynthesis activity, lesser reduction in plant growth and faster recovering ability. Collectively, our results provide strong evidence that AtGolS2 is a useful biotechnological tool to reduce grain yield losses in rice beyond genetic differences under field drought stress.     

栽培稻旱胁迫叶片相关性状的遗传解析

利用籼稻窄叶青8号(ZYQ8)和粳稻京系17(JX17)衍生的加倍单倍体(DH)群体127个株系,2002年在杭州采用田间断水法栽培,在水分胁迫下,对叶片的卷叶、相对含水量和电导率3个性状进行了评价和QTL分析。结果表明,3个性状在DH群体中均存在双向超亲分离,接近正态分布,受数量性状基因的控制;检测到影响这些性状的6个QTL,其中卷叶3个(qLR—1,qLR—5和qLR—11)、相对含水量2个(qRWC—1和qRWC—6和电导率1个(qERC—6）。旱胁迫时,目测卷叶方便易行,适于对大批品种或资源筛选,对抗旱栽培稻品种的筛选和利用具有一定的指导意义。     

Differential responses of antioxidative system to chilling and drought in four rice cultivars differing in sensitivity

Responses of antioxidative defense systems to chilling and drought stresses were comparatively studied in four cultivars of rice (Oryza sativa L.) differing in sensitivity, two of them (Xiangnuo no. 1 and Zimanuo) are tolerant to chilling but sensitive to drought and the other two (Xiangzhongxian no. 2 and IR50) are tolerant to drought but sensitive to chilling. The seedlings of rice were transferred into growth chamber for 5 d at 8 degrees C as chilling treatment, or at 28 degrees C as control, or at 28 degrees C but cultured in 23% PEG-6000 solution as drought stress treatment. Under drought stress the elevated levels of electrolyte leakage, contents of H(2)O(2) and total thiobarbituric acid-reacting substances (TBARS) in Xiangzhongxian no. 2 and IR50 are lower than those in Xiangnuo no. 1 and Zimanuo. On the contrary, Xiangnuo no. 1 and Zimanuo have much lower level of electrolyte leakage, H(2)O(2) and TBARS than Xiangzhongxian no. 2 and IR50 under chilling stress. Activities of antioxidant enzymes (superoxide dismutase (SOD), catalase, and ascorbate-peroxidase (APX)) and contents of antioxidants (ascorbaic acid and reduced glutathione) were measured during the stress treatments. All of them were enhanced greatly until 3 d after drought stress in the two drought-tolerant cultivars, or after chilling stress in the two chilling-tolerant cultivars. They all were decreased at 5 d after stress treatments. On the other hand, activities of antioxidant enzymes and contents of antioxidants were decreased greatly in the drought-sensitive cultivars after drought stress, or in the chilling-sensitive cultivars after chilling stress. The results indicated that tolerance to drought or chilling in rice is well associated with the enhanced capacity of antioxidative system under drought or chilling condition, and that the sensitivity of rice to drought or chilling is linear correlated to the decreased capacity of antioxidative system.     

Small temporal differences in water uptake among varieties of pearl millet (Pennisetum glaucum (L.) R. Br.) are critical for grain yield under terminal drought

Background and aim

Intuitively, access to water from the soil at key phenological stages is important for adaptation to drought. This study aimed to assess the temporal pattern of water extraction under terminal drought stress.

Methods

Pearl millet genotypes with varying levels of terminal drought tolerance were grown in a lysimetric system with a soil volume and plant spacing similar to field conditions. Water extraction was monitored until maturity under differing water regimes.

Results

The yield did not differ among genotypes under well-watered (WW) conditions, and the water extraction profile of WW plants was similar across all genotypes. In contrast, the yield of sensitive genotypes was 30–100 % lower than that of tolerant lines under water stress (WS). The total volumes of water extracted by tolerant and sensitive genotypes were similar under WS; however, tolerant genotypes extracted less water prior to anthesis, and more water after anthesis. Grain yield was positively related to the amount of water extracted during week three after panicle emergence. Increased water extraction after anthesis benefitted the tillers more than the main culm and was correlated with higher staygreen scores.

Conclusion

Increased water uptake after anthesis, which results from earlier water conservation during pre-anthesis, increases yield under terminal drought in pearl millet.     

Comprehensive metabolomic,proteomic and physiological analyses of grain yield reduction in rice under abrupt drought–flood alternation stress

Abrupt drought–flood alternation (T1) is a meteorological disaster that frequently occurs during summer in southern China and the Yangtze river basin, often causing a significant loss of rice production. In this study, the response mechanism of yield decline under abrupt drought–flood alternation stress at the panicle differentiation stage was analyzed by looking at the metabolome, proteome as well as yield and physiological and biochemical indexes. The results showed that drought and flood stress caused a decrease in the yield of rice at the panicle differentiation stage, and abrupt drought–flood alternation stress created a synergistic effect for the reduction of yield. The main reason for the decrease of yield per plant under abrupt drought–flood alternation was the decrease of seed setting rate. Compared with CK0 (no drought and no flood), the net photosynthetic rate and soluble sugar content of T1 decreased significantly and its hydrogen peroxidase, superoxide dismutase, peroxidase activity increased significantly. The identified differential metabolites and differentially expressed proteins indicated that photosynthesis metabolism, energy metabolism pathway and reactive oxygen species response have changed strongly under abrupt drought–flood alteration stress, which are factors that leads to the rice grain yield reduction.