Effects of Favorable Alleles for Water-Soluble Carbohydrates at Grain Filling on Grain Weight under Drought and Heat Stresses in Wheat

Drought, heat and other abiotic stresses during grain filling can result in reductions in grain weight. Conserved water-soluble carbohydrates (WSC) at early grain filling play an important role in partial compensation of reduced carbon supply. A diverse population of 262 historical winter wheat accessions was used in the present study. There were significant correlations between 1000-grain weight (TGW) and four types of WSC, viz. (1) total WSC at the mid-grain filling stage (14 days after flowering) produced by leaves and non-leaf organs; (2) WSC contributed by current leaf assimilation during the mid-grain filling; (3) WSC in non-leaf organs at the mid-grain filling, excluding the current leaf assimilation; and (4) WSC used for respiration and remobilization during the mid-grain filling. Association and favorable allele analyses of 209 genome-wide SSR markers and the four types of WSC were conducted using a mixed linear model. Seven novel favorable WSC alleles exhibited positive individual contributions to TGW, which were verified under 16 environments. Dosage effects of pyramided favorable WSC alleles and significantly linear correlations between the number of favorable WSC alleles and TGW were observed. Our results suggested that pyramiding more favorable WSC alleles was effective for improving both WSC and grain weight in future wheat breeding programs.     

不同生育时期干旱对冬小麦氮素吸收与利用的影响

以抗旱性强的‘石家庄8号’和抗旱性弱的‘偃麦20’冬小麦(Triticum aestivum)为材料, 在田间遮雨棚条件下, 研究返青-拔节期、拔节-开花期和灌浆后期3个生育期不同干旱程度对冬小麦产量、氮素吸收、分配和利用的影响。结果表明, 在干旱条件下, 抗旱性强的‘石家庄8号’产量高于抗旱性弱的‘偃麦20’, 并且其3个生育时期轻度干旱均可提高产量。拔节-开花期干旱对两个冬小麦品种氮素的吸收和运转影响均最大, 其次为返青-拔节期, 而灌浆后期影响较小。不同生育期中度和重度干旱均降低了花前贮藏氮素向籽粒中的转移, 并且氮肥利用效率和生产率也较低, 而在返青-拔节和灌浆后期轻度干旱有利于营养器官的氮素向籽粒中转移, 提高了氮肥利用效率和生产率。在干旱条件下, 抗旱性强的‘石家庄8号’籽粒氮素积累对花前贮藏氮素再运转的依赖程度高, 而‘偃麦20’对花后氮素的积累和转移依赖较高。综合产量和氮素的转移特点, 在生产实践中, 返青-拔节期和灌浆后期要注意对小麦进行适度的干旱处理, 在拔节-开花期要保证冬小麦的充分灌溉, 从而有利于氮素的积累和分配。     

Grain zinc, iron and protein concentrations and zinc-efficiency in wild emmer wheat under contrasting irrigation regimes

Micronutrient malnutrition, and particularly deficiency in zinc (Zn) and iron (Fe), afflicts over three billion people worldwide, and nearly half of the world’s cereal-growing area is affected by soil Zn deficiency. Wild emmer wheat [Triticum turgidum ssp. dicoccoides (Körn.) Thell.], the progenitor of domesticated durum wheat and bread wheat, offers a valuable source of economically important genetic diversity including grain mineral concentrations. Twenty two wild emmer wheat accessions, representing a wide range of drought resistance capacity, as well as two durum wheat cultivars were examined under two contrasting irrigation regimes (well-watered control and water-limited), for grain yield, total biomass production and grain Zn, Fe and protein concentrations. The wild emmer accessions exhibited high genetic diversity for yield and grain Zn, Fe and protein concentrations under both irrigation regimes, with a considerable potential for improvement of the cultivated wheat. Grain Zn, Fe and protein concentrations were positively correlated with one another. Although irrigation regime significantly affected ranking of genotypes, a few wild emmer accessions were identified for their advantage over durum wheat, having consistently higher grain Zn (e.g., 125 mg kg^?1), Fe (85 mg kg^?1) and protein (250 g kg^?1) concentrations and high yield capacity. Plants grown from seeds originated from both irrigation regimes were also examined for Zn efficiency (Zn deficiency tolerance) on a Zn-deficient calcareous soil. Zinc efficiency, expressed as the ratio of shoot dry matter production under Zn deficiency to Zn fertilization, showed large genetic variation among the genotypes tested. The source of seeds from maternal plants grown under both irrigation regimes had very little effect on Zn efficiency. Several wild emmer accessions revealed combination of high Zn efficiency and drought stress resistance. The results indicate high genetic potential of wild emmer wheat to improve grain Zn, Fe and protein concentrations, Zn deficiency tolerance and drought resistance in cultivated wheat.     

干旱调控下小麦RIL群体灌浆期茎秆可溶性碳水化合物积累与转运的遗传分析

以小麦RIL群体(陇鉴19×Q9086,F₈)120个株系及其亲本为供试材料,研究雨养(DS)和正常灌溉(WW)条件下,小麦灌浆期不同发育阶段主茎不同节位可溶性碳水化合物(WSC)含量、转运率及其对籽粒的贡献率,以及穗粒重的遗传特点及各目标性状间的相关性.结果表明: 在两种水分条件下,小麦RIL群体各目标性状变异广泛,变异系数在2.7%～62.1%(DS)和1.9%～52.1%(WW),多样性指数在0.61～0.90(DS)和0.64～0.89(WW),且存在超亲分离.各目标性状表型受基因型、水分环境、节位和发育时期的显著影响.其中,WSC含量受发育时期的影响较大,WSC转运率具有显著的水分和节位主导效应,而WSC转运对籽粒的贡献率受基因型、节位和水分的共同作用.开花初期和灌浆中期WSC含量、花前WSC转运率对籽粒的贡献率之间普遍表现为显著或极显著正相关,且干旱条件下其相关系数更高.各目标性状的遗传力较低(hB₂在干旱条件下为0.31～0.56,灌溉条件下为0.44～0.67),控制各目标性状的遗传基因对数在6～29对(DS)和3～19对(WW).表明该群体对所考察性状有贡献的等位基因在其后代群体中得到广泛分离,其表达易受水分环境的影响,符合典型数量性状特点.     

干旱胁迫下冬小麦光合产物分配格局及其与产量的关系

研究于2005-2006年在中国科学院封丘农田生态试验站防雨棚下进行,在充足供水(W1)、轻度干旱(W2)和严重干旱(W3)条件下,研究冬小麦周麦18(Zhoumai18)和济麦20(Jimai20)干物质积累和分配、可溶性碳水化合物(WSC)含量及其与产量形成的关系。结果显示,随着干旱胁迫程度加重,两个品种冬小麦干物质积累下降,光合产物分配格局改变。干旱胁迫导致花前叶片分配指数下降、茎和叶鞘分配指数上升,花后穗分配指数上升。干旱胁迫下冬小麦拔节期茎WSC减少,但孕穗期、开花期各器官WSC均增加。相关分析表明,冬小麦千粒重与孕穗期、开花期茎和根WSC,与开花期叶片干重显著正相关(P0.05);穗粒数与拔节期茎WSC极显著正相关(P0.01),与开花期茎和叶片干重显著正相关(P0.05)。研究认为拔节期是冬小麦需水关键期,干旱胁迫能促进光合产物向当时生长中心分配,不同生育期茎WSC是冬小麦抗旱育种的重要生理生态指标之一。     

Assessment of transpiration efficiency in peanut (Arachis hypogaea L.) under drought using a lysimetric system

Transpiration efficiency (TE) is an important trait for drought tolerance in peanut ( Arachis hypogaea L.). The variation in TE was assessed gravimetrically using a long time interval in nine peanut genotypes (Chico, ICGS 44, ICGV 00350, ICGV 86015, ICGV 86031, ICGV 91114, JL 24, TAG 24 and TMV 2) grown in lysimeters under well-watered or drought conditions. Transpiration was measured by regularly weighing the lysimeters, in which the soil surface was mulched with a 2-cm layer of polythene beads. TE in the nine genotypes used varied from 1.4 to 2.9 g kg⁻¹ under well-watered and 1.7 to 2.9 g kg⁻¹ under drought conditions, showing consistent variation in TE among genotypes. A higher TE was found in ICGV 86031 in both well-watered and drought conditions and lower TE was found in TAG-24 under both water regimes. Although total water extraction differed little across genotypes, the pattern of water extraction from the soil profile varied among genotypes. High water extraction within 24 days following stress imposition was negatively related to pod yield ( r ²   =   0.36), and negatively related to water extraction during a subsequent period of 32 days ( r ²   =   0.73). By contrast, the latter, i.e. water extraction during a period corresponding to grain filling (24 to 56 days after flowering) was positively related to pod yield ( r ²   =   0.36). TE was positively correlated with pod weight ( r ²   =   0.30) under drought condition. Our data show that under an intermittent drought regime, TE and water extraction from the soil profile during a period corresponding to pod filling were the most important components.     

Selection for water-soluble carbohydrate accumulation and investigation of genetic × environment interactions in an elite wheat breeding population

Key message

Water-soluble carbohydrate accumulation can be selected in wheat breeding programs with consideration of genetic × environmental interactions and relationships with other important characteristics such as relative maturity and nitrogen concentration, although the correlation between WSC traits and grain yield is low and inconsistent.

Abstract

The potential to increase the genetic capacity for water-soluble carbohydrate (WSC) accumulation is an opportunity to improve the drought tolerance capability of rainfed wheat varieties, particularly in environments where terminal drought is a significant constraint to wheat production. A population of elite breeding germplasm was characterized to investigate the potential for selection of improved WSC concentration and total amount in water deficit and well-watered environments. Accumulation of WSC involves complex interactions with other traits and the environment. For both WSC concentration (WSCC) and total WSC per area (WSCA), strong genotype × environment interactions were reflected in the clear grouping of experiments into well-watered and water deficit environment clusters. Genetic correlations between experiments were high within clusters. Heritability for WSCC was larger than for WSCA, and significant associations were observed in both well-watered and water deficit experiment clusters between the WSC traits and nitrogen concentration, tillering, grains per m², and grain size. However, correlations between grain yield and WSCC or WSCA were weak and variable, suggesting that selection for these traits is not a better strategy for improving yield under drought than direct selection for yield.

     

Salinity and drought interaction in wheat (Triticum aestivum L.) is affected by the genotype and plant growth stage

Salinity and drought are important agro-environmental problems occurring separately as well as together with the combined occurrence increasing with time due to climate change. Screening of bread wheat genotypes against salinity or drought alone is common; however, little information is available on the response of wheat genotypes to a combination of these stresses. This study investigates the response of a salt-resistant (SARC-1) and a salt-sensitive (7-Cerros) wheat genotype to drought at different growth stages under non-saline (ECe 2.1 dS m^?1) and saline soil (ECe 15 dS m^?1) conditions. Drought was applied by withholding water for 21 days at a particular growth stage viz. tillering, booting, and grain filling stages. At booting stage measurements regarding water relations, leaf ionic composition and photosynthetic attributes were made. At maturity grain yield and different yield, components were recorded. Salinity and drought significantly decreased grain yield and different yield components with a higher decrease in the case of combined stress of salinity × drought. The complete drought treatment (drought at tillering + booting + grain filling stages) was most harmful for wheat followed by drought at booting stage and grain filling–tillering stages, respectively. The salt-resistant wheat genotype SARC-1 performed better than the salt-sensitive genotype 7-Cerros in different stress treatments. A decrease in the water and turgor potentials, photosynthetic and transpiration rates, stomatal conductance, leaf K⁺, and increased leaf Na⁺ were the apparent causes of growth and yield reduction of bread wheat due to salinity, drought, and salinity × drought.     

Comparison of fructan dynamics in two wheat cultivars with different capacities of accumulation and remobilization under drought stress

Remobilization of stored carbohydrates in the stem of wheat plants is an important contributor to grain filling under drought stress (DS) conditions. A massive screening on Iranian wheat cultivars was performed based on stem dry weight changes under well-watered and DS conditions. Two cultivars, Shole and Crossed Falat Hamun (CFH), with different fructan accumulation and remobilization behavior were selected for further studies. Water-soluble carbohydrates (WSCs) and fructan metabolizing enzymes were studied both in the stem penultimate and in sucrose (Suc) treated, excised leaves. Under drought, CFH produced higher grain yields than Shole (412 vs 220 g m(-2)). Also, grain yield loss under drought was more limited in CFH than in Shole (17 vs 54%). Under drought, CFH accumulated more graminan-type fructo-oligosaccharides than Shole. After anthesis, fructan 6-exohydrolase (6-FEH; EC 3.2.1.154) activities increased more prominently than fructan 1-exohydrolase (EC 3.2.1.153) activities during carbon remobilization. Interestingly, CFH showed higher 6-FEH activities in the penultimate than Shole. The field experiment results suggest that the combined higher remobilization efficiency and high 6-FEH activities in stems of wheat could contribute to grain yield under terminal drought. Similar to the penultimate, fructan metabolism differed strongly in Suc-treated detached leaves of selected cultivars. This suggests that variation in the stem fructan among wheat cultivars grown in the field could be traced by leaf blade induction experiments.     

Effects of drought stress and selenium supply on growth and physiological characteristics of wheat seedlings

The paper studied the effects of drought stress, selenium (Se) supply and their combination on growth and physiological characteristics of wheat (Triticum aestivum L. cv Shijiazhuang NO. 8) seedlings. The experimental design included two water treatments (well-watered, 75% of maximum field capacity; drought stress, 30% of maximum field capacity) and two Se levels (0; 0.5 mg/kg) to determine whether Se can modify the negative impacts of drought stress on seedling growth and physiological traits. Drought stress caused a marked decline in growth parameters and soluble protein content, whereas it induced an increase in root activity, proline content and the activities of peroxidase (POD) and catalase (CAT) of leaf tissue. On the other hand, Se supply induced an increase in biomass accumulation only under well-watered condition. Under drought stress, Se supply increased free proline content, root activity and the activities of POD and CAT in leaf tissue, but did not significantly affect on growth parameters. These results implied that drought stress brought harmful effects on wheat seedlings, and that Se supply was favorable for biomass accumulation of wheat seedlings under well-watered condition. However, it did not significantly affect on biomass accumulation under drought stress, although it increased root activity and activities of some antioxidant index in experimental periods.     

Association mapping of dynamic developmental plant height in common wheat

Drought as a major abiotic stress often occurs from stem elongation to the grain filling stage of wheat in northern China. Plant height (PH) is a suitable trait to model the dissection of drought tolerance. The purposes of the present study were to validate molecular markers for PH developmental behavior and identify elite alleles of molecular markers. After the phenotyping of 154 accessions for PH dynamic development under well-watered (WW) and drought stressed (DS) conditions, and the genotyping of 60 SSR markers from six candidate chromosome regions related to PH found in our previous linkage mapping studies, both parameters PH and drought tolerance coefficient (DTC) calculated by the conditional analysis were used for association mapping. A total of 46 significant association signals (P < 0.01) were identified in 23 markers, and phenotypic variation ranged from 7 to 50%. Among them, four markers Xgwm261-2D, Xgwm495-4B, Xbarc109-4B and Xcfd23-4D were detected under both water regimes. Furthermore, 10 markers were associated with DTC, and four with both parameters PH and DTC at the same plant development stage. The results revealed different allelic effects of associated markers; for example, the 155 bp Xgwm495-4B allele was associated with a reduced height of −11.2 cm under DS and −15.3 cm under WW, whereas the 167 bp allele exhibited increased height effects of 3.9 and 8.1 cm, respectively. This study demonstrates a strong power of joint association analysis and linkage mapping for the identification of important genes in wheat.     

灌溉对干旱区冬小麦干物质积累、分配和产量的影响

为探明灌溉对干旱区冬小麦(Triticum aestivum)产量、水分利用效率(WUE)、干物质积累及分配等的影响, 以甘肃河西走廊冬小麦适宜种植品种‘临抗2号’为材料进行了研究。在冬季灌水180 mm的条件下, 生育期以灌水量和灌水次数等共设置5个处理, 分别为: 拔节期灌水量165 mm (W1)、拔节期灌水量120 mm +抽穗期灌水量105 mm (W2)、拔节期灌水量105 mm +抽穗期灌水量105 mm +灌浆期灌水量105 mm (W3)、拔节期灌水量75 mm +抽穗期灌水量75 mm +灌浆期灌水量75 mm (W4)、拔节期灌水量105 mm +抽穗期灌水量75 mm +灌浆期灌水量45 mm (W5)。结果表明: 随着生育期的推进, 土壤有效含水量(AWC)受灌水次数及灌水量影响更加明显; W3、W4处理的土壤各层AWC在灌浆期均较高; 叶面积指数(LAI)下降慢, 延缓了生育后期的衰老; 生育后期干物质积累增加, 提高了穗粒数、千粒重和籽粒产量。籽粒产量以W3处理最高, 但W4具有最高的WUE, 且籽粒产量与W3无显著差异, 但W4较灌溉总量相同的W2和W5以及灌水量最少的W1具有明显的指标优势。W1、W2、W5处理灌浆期各层土壤AWC均较低, 花后LAI下降快, 干物质积累减少, 灌浆持续期缩短, 穗粒数和千粒重减少, 最终表现为籽粒产量和WUE下降。灌浆期水分胁迫可促进花前储存碳库向籽粒的再转运, 并随着干旱胁迫的加重而提高, 对籽粒产量起补偿作用; 水分胁迫提高了灌浆速率, 但缩短了灌浆持续期。相关性分析表明, 灌浆持续期、有效灌浆持续期、有效灌浆期粒重增加值和最大籽粒灌浆速率出现时间与千粒重和籽粒产量均呈正相关。综合考虑, 拔节、抽穗及灌浆期各灌溉75 mm是高产高WUE的最佳灌水方案。     

Foliar application of zinc improves morpho-physiological and antioxidant defense mechanisms,and agronomic grain biofortification of wheat (Triticum aestivum L.) under water stress

Agronomic biofortification with zinc (Zn) may be engaged to improve the nutritious value of food crops along-with tolerance to water deficit conditions. The Zn may increase plant resistance to water stress by boosting physiological and enzymatic antioxidants defense mechanisms. Major objective of this study was to investigate the effect of foliar applied Zn on grain zin biofortification and drought tolerance in wheat. Treatments include application of Zinc at terminal growth phases (BBCH growth stage 49 and BBCH growth stage 65) with five levels: 0 (control-ck), water spray, 5, 10 and 15 mM under two levels of water regimes; well-watered (where 80% water holding capacity (WHC) was maintained in the soil) and water stress, (where 40% WHC was maintained in the soil). Results revealed that water stress significantly reduced relative water contents, gas exchange attributes, plant height, yield and yield related attributes of wheat. In contrast, hydrogen peroxide, free proline levels, activities of malondialdehyde, and concentration of soluble protein were markedly increased under water stress condition. Application of various levels of Zn significantly improved the CAT, SOD, POD and ASP activities at 40% WHC compared with control treatment. Foliarly applied 10 and 15 mM Zn predominantly reduced the damaging impact of water stress by improving the plant status in the form of plant height, RWC and gas exchange attributes. Likewise, wheat plant treated with 10 mM Zn under water stress condition increased the grain yield by improving number of grains per spike, 100 grain weight and biological yield compared with control. Moreover, increasing Zn levels also increased Zn concentration in grains and leaves. Overall, this study suggests that optimum level of Zn (10 mM) might be promising for alleviating the adverse impacts of water stress and enhance the grain biofortification in wheat.     

Terminal drought and seed priming improves drought tolerance in wheat

Plants retain the preceding abiotic stress memory that may aid in attainment of tolerance to subsequent stresses. This study was conducted to evaluate the influence of terminal drought memory (drought priming) and seed priming in improving drought tolerance in wheat (Triticum aestivum L.). During first growing season, wheat was planted in field under optimal (well-watered) and drought stress imposed at reproductive stage (BBCH growth stage 49) until maturity (BBCH growth stage 83). Seeds collected from both sources were subjected to hydropriming or osmopriming (with 1.5% CaCl₂ solution); while, dry seed was taken as control. Treated and control seeds, from both sources, were sown in soil filled pots. After the completion of seedling emergence, pots were maintained at 50% water holding capacity (drought) or 100% water holding capacity (well-watered). Drought stress suppressed the plant growth (2–44%), perturbed water relations (1–18%) and reduced yield (192%); however, osmolytes accumulation (3–14%) and malondialdehyde contents (26–29%) were increased under drought. The crop raised from the seeds collected from terminal drought stressed plants had better growth (5–63%), improved osmolyte accumulation (13–45%), and lower lipid peroxidation (3%) than the progeny of well-watered crop. Seed priming significantly improved the crop performance under drought stress as compared to control. However, osmopriming was more effective than hydropriming in this regard as it improved leaf area (9–43%), tissue water status (2–47%), osmolytes accumulation (6–48%) and grain yield (14–79%). In conclusion, terminal drought induced modifications in seed composition and seed priming improved transgenerational drought tolerance through improvement in tissue water status and osmolytes accumulation, and decrease in lipid peroxidation.     

Expression analysis of the genes involved in accumulation and remobilization of assimilates in wheat stem under terminal drought stress

Stem reserve mobilization and expression of major genes involved in fructan metabolism during grain filling in wheat (Triticum aestivum L.) cultivars, Zagros and Marvdasht were studied under terminal drought through withholding water at the anthesis. Mobilized dry matter, maximum specific weight and mobilization efficiency were observed to be higher in the internodes of tolerant cultivar (Zagros), both under well-watered and stress conditions, which resulted in enhanced translocation of stem reserves to the grains. Water soluble carbohydrates (WSC) and its constituent compounds were observed to be higher in the internodes of Zagros than those of sensitive cultivar (Marvdasht). Maximum relative expression of 1-SST, 6-SFT, INV, 1-FEHw1, 1-FEHw2, 1-FEHw3 and 6-FEH was significantly higher in the peduncle and penultimate of Zagros compared to Marvdasht cultivar under both drought and well-watered conditions. Expression of 1-FEHw3 and 6-FEH were increased during carbon remobilization in Zagros cultivar, suggesting that both genes are necessary for an efficient degradation and translocation of stem fructans. The mRNA levels of two fructan synthetic enzymes (1-SST and 6-SFT) in the stem were positively correlated with stem WSC concentrations, while the mRNA levels of enzymes involved in fructan hydrolysis (INV, 1-FEHw3 and 6-FEH) were inversely correlated with WSC concentration. According to the achieved results, it can be concluded that certain characteristics of Zagros cultivar, enhanced capability of fructan storage, higher mobilization efficiency and high gene expression level of 1-SST, 6-SFT, 1-FEHw3 as well as 6-FEH genes might help the drought tolerant cultivar to cope the stress conditions.     

Differences in Sugar Accumulation and Mobilization between Sequential and Non-Sequential Senescence Wheat Cultivars under Natural and Drought Conditions

Wheat leaf non-sequential senescence at the late grain-filling stage involves the early senescence of younger flag leaves compared to that observed in older second leaves. On the other hand, sequential senescence involves leaf senescence that follows an age-related pattern, in which flag leaves are the latest to undergo senescence. The characteristics of sugar metabolism in two sequential senescence cultivars and two non-sequential senescence cultivars under both natural and drought conditions were studied to elucidate the underlying mechanism of drought tolerance in two different senescence modes. The results showed that compared to sequential senescence wheat cultivars, under natural and drought conditions, non-sequential senescence wheat cultivars showed a higher leaf net photosynthetic rate, higher soluble sugar levels in leaves, leaf sheaths, and internodes, higher leaf sucrose synthase (SS) and sucrose phosphate synthase (SPS) activity, and higher grain SS activity, thereby suggesting that non-sequential senescence wheat cultivars had stronger source activity. Spike weight, grain weight per spike, and 100-grain weight of non-sequential senescence cultivars at maturity were significantly higher than those of sequential senescence cultivars under both natural and drought conditions. These findings indicate that the higher rate of accumulation and the higher mobilization of soluble sugar in the leaves, leaf sheaths and internodes of non-sequential senescence cultivars improve grain weight and drought tolerance. At the late grain-filling stage, drought conditions adversely affected leaf chlorophyll content, net photosynthetic rate, soluble sugar and sucrose content, SS and SPS activity, gain SS activity, and weight. This study showed that higher rates of soluble sugar accumulation in the source was one of the reasons of triggering leaf non-sequential senescence, and higher rates of soluble sugar mobilization during leaf non-sequential senescence promoted high and stable wheat yield and drought tolerance.     

Screening for Drought Resistance of Rice Recombinant Inbred Populations in the Field

In a 2-year experiment, 187 genotypes were grown under well-watered and drought stress conditions, imposed at panicle initiation stage. The relationship of genotypic variation in yield under drought conditions to potential yield, heading date and flowering delay, reduction in plant height, and to a drought response index (DRI) was detected. Grain yield under drought stress conditions was associated with yield under well-watered conditions (r= 0.47**, and r= 0.61** during 2 years of tests). The delay of heading date ranged from -1 (no delay) to 24days, and was negatively associated with grain yield(r=-0.40*), spikelet fertility percentage (r=-0.40**), harvest index (r=-0.58**), but positively associated with yield reduction percentage (r= 0.60**). The reduction in plant height was negatively associated with grain yield (r =-0.24**, and r =-0.29**), spikelet fertility percentage (r =-0.23**, and r =-0.21*), harvest index (r =-0.37**, and r = -0.54**), and positively associated with yield reduction percentage (r = 0.58**, and r = 0.58**) in 2003 and 2004, respectively. The DRI of genotypes was strongly associated with grain yield (r = 0.87**, and r= 0.77**), fertility percentage (r= 0.66** and r = 0.54**), harvest index (r=0.67** and r=0.61**), and negatively associated with grain reduction percentage (r=-0.70**, and r=-0.73**)under drought stress. The results indicate that genotypes with drought resistance can be identified by measuring yield potential, delay in flowering, reduction in plant height, or DRI under test environments of well-watered and drought stress.