Mapping of post-flowering drought resistance traits in grain sorghum: association between QTLs influencing premature senescence and maturity

The identification of genetic factors underlying the complex responses of plants to drought stress provides a solid basis for improving drought resistance. The stay-green character in sorghum (Sorghum bicolor L. Moench) is a post-flowering drought resistance trait, which makes plants resistant to premature senescence under drought stress during the grainfilling stage. The objective of this study was to identify quantitative trait loci (QTLs) that control premature senescence and maturity traits, and to investigate their association under post-flowering drought stress in grain sorghum. A genetic linkage map was developed using a set of recombinant inbred lines (RILs) obtained from the cross B35 × Tx430, which were scored for 142 restriction fragment length polymorphism (RFLP) markers. The RILs and their parental lines were evaluated for post-flowering drought resistance and maturity in four environments. Simple interval mapping identified seven stay-green QTLs and two maturity QTLs. Three major stay-green QTLs (SGA, SGD and SGG) contributed to 42% of the phenotypic variability (LOD 9.0) and four minor QTLs (SGB, SGI.1, SGI.2, and SGJ) significantly contributed to an additional 25% of the phenotypic variability in stay-green ratings. One maturity QTL (DFB) alone contributed to 40% of the phenotypic variability (LOD 10.0), while the second QTL (DFG) significantly contributed to an additional 17% of the phenotypic variability (LOD 4.9). Composite interval mapping confirmed the above results with an additional analysis of the QTL × Environment interaction. With heritability estimates of 0.72 for stay-green and 0.90 for maturity, the identified QTLs explained about 90% and 63% of genetic variability for stay-green and maturity traits, respectively. Although stay-green ratings were significantly correlated (r=0.22, P ≤ 0.05) with maturity, six of the seven stay-green QTLs were independent of the QTLs influencing maturity. Similarly, one maturity QTL (DFB) was independent of the stay-green QTLs. One stay-green QTL (SGG), however, mapped in the vicinity of a maturity QTL (DFG), and all markers in the vicinity of the independent maturity QTL (DFB) were significantly (P ≤ 0.1) correlated with stay-green ratings, confounding the phenotyping of stay-green. The molecular genetic analysis of the QTLs influencing stay-green and maturity, together with the association between these two inversely related traits, provides a basis for further study of the underlying physiological mechanisms and demonstrates the possibility of improving drought resistance in plants by pyramiding the favorable QTLs. Received: 10 October 1998 / Accepted: 12 July 1999     

Mapping QTLs associated with drought resistance in sorghum (Sorghum bicolor L. Moench)

Drought is a major abiotic stress factor limiting crop production. Identification of genetic factors involved in plant responses to drought stress will provide a solid foundation to improve drought resistance. Sorghum is well adapted to hot dry environments and regarded as a model for studying drought resistance among the grasses. Significant progress in genome mapping of this crop has also been made. In sorghum, rapid premature leaf death generally occurs when water is limited during the grain filling period. Premature leaf senescence, in turn, leads to charcoal rot, stalk lodging, and significant yield loss. More than 80% of commercial sorghum hybrids in the United States are grown under non-irrigated conditions and although most of them have pre-flowering drought resistance, many do not have any significant post-flowering drought resistance. Stay-green is one form of drought resistance mechanism, which gives sorghum resistance to premature senescence under soil moisture stress during the post-flowering period. Quantitative trait locus (QTL) studies with recombinant inbred lines (RILs) and near-isogenic lines (NILs) identified several genomic regions associated with resistance to pre-flowering and post-flowering drought stress. We have identified four genomic regions associated with the stay-green trait using a RIL population developed from B35 × Tx7000. These four major stay-green QTLs were consistently identified in all field trials and accounted for 53.5% of the phenotypic variance. We review the progress in mapping stay-green QTLs as a component of drought resistance in sorghum. The molecular genetic dissection of the QTLs affecting stay-green will provide further opportunities to elucidate the underlying physiological mechanisms involved in drought resistance in sorghum and other grasses.     

Sorghum stay-green QTL individually reduce post-flowering drought-induced leaf senescence

Sorghum is an important source of food, feed, and biofuel, especially in the semi-arid tropics because this cereal is well adapted to harsh, drought-prone environments. Post-flowering drought adaptation in sorghum is associated with the stay-green phenotype. Alleles that contribute to this complex trait have been mapped to four major QTL, Stg1-Stg4, using a population derived from BTx642 and RTx7000. Near-isogenic RTx7000 lines containing BTx642 DNA spanning one or more of the four stay-green QTL were constructed. The size and location of BTx642 DNA regions in each RTx7000 NIL were analysed using 62 DNA markers spanning the four stay-green QTL. RTx7000 NILs were identified that contained BTx642 DNA completely or partially spanning Stg1, Stg2, Stg3, or Stg4. NILs were also identified that contained sub-portions of each QTL and various combinations of the four major stay-green QTL. Physiological analysis of four RTx7000 NILs containing only Stg1, Stg2, Stg3, or Stg4 showed that BTx642 alleles in each of these loci could contribute to the stay-green phenotype. RTx7000 NILs containing BTx642 DNA corresponding to Stg2 retained more green leaf area at maturity under terminal drought conditions than RTx7000 or the other RTx7000 NILs. Under post-anthesis water deficit, a trend for delayed onset of leaf senescence compared with RTx7000 was also exhibited by the Stg2, Stg3, and Stg4 NILs, while significantly lower rates of leaf senescence in relation to RTx7000 were displayed by all of the Stg NILs to varying degrees, but particularly by the Stg2 NIL. Greener leaves at anthesis relative to RTx7000, indicated by higher SPAD values, were exhibited by the Stg1 and Stg4 NILs. The RTx7000 NILs created in this study provide the starting point for in-depth analysis of stay-green physiology, interaction among stay-green QTL and map-based cloning of the genes that underlie this trait.     

冬小麦叶片持绿能力及其衰老特征研究

以12个冬小麦(Triticum aestivum L.)品种为供试材料,通过田间实验连续2年于开花后定期测定各品种的绿叶数目、绿叶面积、叶绿素和MDA含量以及SOD和CAT活性等指标,并以生理成熟时的保绿度、衰老启动时间为指标进行Hierarchical聚类分析,对小麦品种持绿能力进行分级.结果表明,参试冬小麦品种可分为持绿和非持绿两种类型,‘潍麦8号'(WM8)和‘豫麦66'(YM66)两年均表现为持绿型小麦.在整个灌浆期,持绿型小麦品种绿叶数目、面积、叶绿素含量明显高于非持绿型品种,叶片保护酶SOD与CAT活性也较非持绿小麦强,而其MDA含量明显低于非持绿型小麦品种.持绿型小麦叶片衰老启动时间延迟,生育后期绿叶面积较大,光合作用时间延长,具有较高的产量.本研究结果为冬小麦的品种选育、布局等相关研究奠定基础.     

Stay-green trait serves as yield stability attribute under combined heat and drought stress in wheat (Triticum aestivum L.)

Plant Growth Regulation - The present study was conducted to evaluate the effect of a stay-green trait to stabilize the yield responses under combined heat and drought stress on wheat. Ten...     

Terminal Heat Tolerance in Bread Wheat Determined by Agronomical Traits and SSR Markers

Since global warming affects wheat cropping systems, more has yet to be indicated on the parameters, which control terminal heat tolerance, and severely influence wheat (Triticum aestivum L.) productivity. Identification of tolerant wheat genotypes by heat tolerance-linked molecular markers is a rapid and cost-effective screening tool in plant breeding. Accordingly, in a four-year field experiment (2015–2019), 44 wheat genotypes were selected out of 100 genotypes, and were examined in timely and late planting (mid-January resulting in heat stress). Stress decreased yield components, including 1000-kernel weight (TKW), grains per spike, and plants per square meter, and the physiological traits, including days to heading and days to maturity, grain filling duration, and greenness, and eventually decreased grain yield up to?~?28%. The early maturity genotypes resulted in higher yields under stress conditions by a stress-avoidance mechanism. Among 14 SSR markers, GWM577 was positively correlated with yield, and WMS3062, GWM261, and WMS1025 had positive correlations with longevity under stress. Accordingly, WMS3062 and GWM261 can be used to determine high yield and early maturity genotypes. Furthermore, GWM114 showed a positive correlation with TKW, indicating their usefulness for grouping wheat genotypes and for identifying heat-related markers. Since the crossing of the genetically distant genotypes can create more diverse populations, the results could be applied to plan breeding projects to establish more diverse populations for different chromosomal locations and traits under heat stress conditions. Moreover, our findings demonstrated that the morphological and molecular analyses could be useful for describing wheat genetic variation of heat tolerance.

     

Improved drought resistance in a wheat stay-green mutant tasg1 under field conditions

We investigated the drought resistance of a wheat (Triticum aestivum L.) stay-green mutant tasg1 and its wild-type (WT) in field experiments conducted for two years. Drought stress was imposed by controlling irrigation and sheltering the plants from rain. Compared with the WT, tasg1 exhibited a distinct delayed senescence under both normal and drought stress conditions, as indicated by slower degradation of chlorophyll and decrease in net photosynthetic rate than in WT. At the same time, tasg1 mutants maintained more integrated chloroplasts and thylakoid ultrastructure than did WT plants under drought stress. Lower malondialdehyde content and higher antioxidative enzyme activities in tasg1, compared to WT, may be involved in the stay-green phenotype and drought resistance of tasg1.     

Identification of Differentially Senescing Mutants of Wheat and Impacts on Yield,Biomass and Nitrogen Partitioning

Increasing photosynthetic capacity by extending canopy longevity during grain filling using slow senescing stay-green genotypes is a possible means to improve yield in wheat.Ethyl methanesulfonate (EMS...     

Genetic Architecture of Delayed Senescence,Biomass, and Grain Yield under Drought Stress in Cowpea

The stay-green phenomenon is a key plant trait with wide usage in managing crop production under limited water conditions. This trait enhances delayed senescence, biomass, and grain yield under drought stress. In this study we sought to identify QTLs in cowpea (Vigna unguiculata) consistent across experiments conducted in Burkina Faso, Nigeria, Senegal, and the United States of America under limited water conditions. A panel of 383 diverse cowpea accessions and a recombinant inbred line population (RIL) were SNP genotyped using an Illumina 1536 GoldenGate assay. Phenotypic data from thirteen experiments conducted across the four countries were used to identify SNP-trait associations based on linkage disequilibrium association mapping, with bi-parental QTL mapping as a complementary strategy. We identified seven loci, five of which exhibited evidence suggesting pleiotropic effects (stay-green) between delayed senescence, biomass, and grain yield. Further, we provide evidence suggesting the existence of positive pleiotropy in cowpea based on positively correlated mean phenotypic values (0.34< r <0.87) and allele effects (0.07< r <0.86) for delayed senescence and grain yield across three African environments. Three of the five putative stay-green QTLs, Dro-1, 3, and 7 were identified in both RILs and diverse germplasm with resolutions of 3.2 cM or less for each of the three loci, suggesting that these may be valuable targets for marker-assisted breeding in cowpea. Also, the co-location of early vegetative delayed senescence with biomass and grain yield QTLs suggests the possibility of using delayed senescence at the seedling stage as a rapid screening tool for post-flowering drought tolerance in cowpea breeding. BLAST analysis using EST sequences harboring SNPs with the highest associations provided a genomic context for loci identified in this study in closely related common bean (Phaseolus vulgaris) and soybean (Glycine max) reference genomes.     

Genomic dissection of drought resistance in durum wheat × wild emmer wheat recombinant inbreed line population

Drought is the major factor limiting wheat productivity worldwide. The gene pool of wild emmer wheat, Triticum turgidum ssp. dicoccoides , harbours a rich allelic repertoire for morpho-physiological traits conferring drought resistance. The genetic and physiological bases of drought responses were studied here in a tetraploid wheat population of 152 recombinant inbreed lines (RILs), derived from a cross between durum wheat (cv. Langdon) and wild emmer (acc# G18-16), under contrasting water availabilities. Wide genetic variation was found among RILs for all studied traits. A total of 110 quantitative trait loci (QTLs) were mapped for 11 traits, with LOD score range of 3.0–35.4. Several QTLs showed environmental specificity, accounting for productivity and related traits under water-limited (20 QTLs) or well-watered conditions (15 QTLs), and in terms of drought susceptibility index (22 QTLs). Major genomic regions controlling productivity and related traits were identified on chromosomes 2B, 4A, 5A and 7B. QTLs for productivity were associated with QTLs for drought-adaptive traits, suggesting the involvement of several strategies in wheat adaptation to drought stress. Fifteen pairs of QTLs for the same trait were mapped to seemingly homoeologous positions, reflecting synteny between the A and B genomes. The identified QTLs may facilitate the use of wild alleles for improvement of drought resistance in elite wheat cultivars.     

Gene action of canopy temperature in bread wheat under diverse environments

While canopy temperature (CT) shows a strong and reliable association with yield under drought and heat stress and is used in wheat breeding to select for yield, little is known of its genetic control. The objective of this study was to determine the gene action controlling CT in five wheat populations grown in diverse environments (heat, drought, and well-irrigated conditions). CT showed negative phenotypic correlations with grain yield under drought and well-irrigated environments. Additive × additive effects were most prevalent and significant for all crosses and environments. Dominance and dominance × dominance gene actions were also found, though the significance and direction was specific for each environment and genotypic cross. The use of CT as a selection criterion to improve tolerance to drought was supported by its significant association with grain yield and the genotype differences observed between cultivars. Our results indicated that genetic gains for CT in wheat could be achieved through conventional breeding. However, given some dominance and epistatic effects, it would be necessary to delay the selection process until the frequency of heterozygous loci within families is reduced.     

QTL for yield and associated traits in the Seri/Babax population grown across several environments in Mexico, in the West Asia, North Africa, and South Asia regions

Heat and drought adaptive quantitative trait loci (QTL) in a spring bread wheat population resulting from the Seri/Babax cross designed to minimize confounding agronomic traits have been identified previously in trials conducted in Mexico. The same population was grown across a wide range of environments where heat and drought stress are naturally experienced including environments in Mexico, West Asia, North Africa (WANA), and South Asia regions. A molecular genetic linkage map including 475 marker loci associated to 29 linkage groups was used for QTL analysis of yield, days to heading (DH) and to maturity (DM), grain number (GM2), thousand kernel weight (TKW), plant height (PH), canopy temperature at the vegetative and grain filling stages (CTvg and CTgf), and early ground cover. A QTL for yield on chromosome 4A was confirmed across several environments, in subsets of lines with uniform allelic expression of a major phenology QTL, but not independently from PH. With terminal stress, TKW QTL was linked or pleiotropic to DH and DM. The link between phenology and TKW suggested that early maturity would favor the post—anthesis grain growth periods resulting in increased grain size and yields under terminal stress. GM2 and TKW were partially associated with markers at different positions suggesting different genetic regulation and room for improvement of both traits. Prediction accuracy of yield was improved by 5 % when using marker scores of component traits (GM2 and DH) together with yield in multiple regression. This procedure may provide accumulation of more favorable alleles during selection.     

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.     

Modeling and mapping QTL for senescence-related traits in winter wheat under high temperature

Senescence is a genetically programmed and environmentally influenced process resulting in the destruction of chlorophyll and remobilization of nutrients to younger or reproductive parts of plants. Delayed senescence, or stay-green, contributes to a long grain-filling period and stable yield under stress. To model senescence and identify quantitative trait loci (QTL) for the trait, a population of recombinant inbred lines (RIL) from a cross between winter wheat cultivars, ‘Ventnor’ and ‘Karl 92’ was evaluated for heat tolerance under optimum temperature of 20/15°C (day/night) and continuous heat stress of 30/25°C from 10 days after anthesis (DAA) until maturity. Ventnor is a heat-tolerant cultivar and Karl 92 is a relatively heat-susceptible cultivar. Green leaf area was measured and used to model percent greenness retained over the reproductive period. Chlorophyll content and chlorophyll fluorescence were recorded on flag leaves. Senescence was converted to a quantitative trait using the model. Based on the modeled parameters, the RILs were categorized into three groups. When senescence-related traits were evaluated, nine QTL for heat tolerance were found on chromosome 2A, two each on chromosomes 6A and 6B and one each on chromosome 3A, 3B, and 7A. Both parents contributed favorable alleles for most of the senescence-related traits. Microsatellite markers Xgwm356 and Xgwm5 prominently linked to the senescence-related traits may be useful in marker-assisted breeding. These and the linked AFLP (amplified fragment length polymorphism) markers XCGT.TGCG-349, XCGT.GTG-343, and XCGT.CTCG-406, if converted to STS (sequence tagged sites), can be used for further molecular dissection of the QTL for post-anthesis heat tolerance.     

Physiological and morphological traits associated with adaptation of lucerne (Medicago sativa) to severely drought‐stressed and to irrigated environments

Adaptation to severe drought and to irrigated cropping can both contribute to increased water use efficiency of lucerne, but knowledge on the relevant adaptive traits is limited. Five cultivars featuring contrasting adaptive responses for 3‐year forage yield across 10 agricultural environments of the western Mediterranean basin were currently studied, to identify physiological and morphological traits associated with specific and wide‐adaptation responses. The landraces Mamuntanas, Demnat 203 and Erfoud 1, and the varieties SARDI 10 and Prosementi, were grown in replicated metal containers (55 cm long × 12 cm wide × 75 cm deep; 21 plants per container) under irrigation (weekly restoring soil field capacity) and under moderate and severe drought stress (implying decreased irrigation for 30 days followed by withheld irrigation for 33 and 58 days, respectively). Cultivar post‐stress survival reflected the known cultivar adaptation to drought‐prone agricultural environments. Demnat 203, specifically adapted to irrigated, frequently mown environments, displayed higher amounts of starch, soluble proteins and total nitrogen in the crown and the root under irrigation. This was due to outstanding organ size and, for starch, higher concentrations. Mamuntanas, specifically adapted to drought‐prone environments, exhibited high water‐soluble carbohydrate concentration in storage organs under severe stress, along with a water‐conservation strategy implying less water used in initial drought‐stress phases due to limited early root development that resulted in more water available under severe stress. Drought‐tolerant germplasm also displayed lower wilting under early stress, more plants with green tissues under severe stress, and more stems per plant in stress or favourable conditions. Multivariate patterns of cultivar variation for physiological and morphological traits were strictly associated with cultivar variation for adaptation pattern. Our results highlighted the difficulty to combine some traits of environment‐specific adaptive value into a unique widely adapted variety, supporting the selection of varieties specifically adapted to irrigated or severely drought‐prone environments.     

QTL for nodal root angle in sorghum (<Emphasis Type="Italic">Sorghum bicolor</Emphasis> L. Moench) co-locate with QTL for traits associated with drought adaptation

Nodal root angle in sorghum influences vertical and horizontal root distribution in the soil profile and is thus relevant to drought adaptation. In this study, we report for the first time on the mapping of four QTL for nodal root angle (qRA) in sorghum, in addition to three QTL for root dry weight, two for shoot dry weight, and three for plant leaf area. Phenotyping was done at the six leaf stage for a mapping population (n = 141) developed by crossing two inbred sorghum lines with contrasting root angle. Nodal root angle QTL explained 58.2% of the phenotypic variance and were validated across a range of diverse inbred lines. Three of the four nodal root angle QTL showed homology to previously identified root angle QTL in rice and maize, whereas all four QTL co-located with previously identified QTL for stay-green in sorghum. A putative association between nodal root angle QTL and grain yield was identified through single marker analysis on field testing data from a subset of the mapping population grown in hybrid combination with three different tester lines. Furthermore, a putative association between nodal root angle QTL and stay-green was identified using data sets from selected sorghum nested association mapping populations segregating for root angle. The identification of nodal root angle QTL presents new opportunities for improving drought adaptation mechanisms via molecular breeding to manipulate a trait for which selection has previously been very difficult.     

Dissecting Maize Productivity: Ideotypes Associated with Grain Yield under Drought Stress and Well-watered Conditions

To increase maize (Zea mays L.) yields in drought‐prone environments and offset predicted maize yield losses under future climates, the development of improved breeding pipelines using a multi‐disciplinary approach is essential. Elucidating key growth processes will provide opportunities to improve drought breeding progress through the identification of key phenotypic traits, ideotypes, and donors. In this study, we tested a large set of tropical and subtropical maize inbreds and single cross hybrids under reproductive stage drought stress and well‐watered conditions. Patterns of biomass production, senescence, and plant water status were measured throughout the crop cycle. Under drought stress, early biomass production prior to anthesis was important for inbred yield, while delayed senescence was important for hybrid yield. Under well‐watered conditions, the ability to maintain a high biomass throughout the growing cycle was crucial for inbred yield, while a stay‐green pattern was important for hybrid yield. While new quantitative phenotyping tools such as spectral reflectance (Normalized Difference Vegetation Index, NDVI) allowed for the characterization of growth and senescence patterns as well as yield, qualitative measurements of canopy senescence were also found to be associated with grain yield.     

高CO2浓度、干旱及其互作对不同持绿型小麦幼苗的影响

以小麦持绿型品种‘烟农19'和非持绿型品种‘旱选3号'为试材,在开放式气室(OTC)内设置正常CO₂浓度(370 μmol·mol^-1)和高CO₂浓度(550 μmol·mol^-1),干旱(田间持水量的45%～55%)和灌溉(田间持水量的75%～85%),共4个环境处理,采用盆栽法研究高CO₂浓度、干旱及其互作对不同持绿型小麦幼苗生长性状、生物量积累和生理性状的影响。结果表明: 干旱显著抑制了小麦幼苗的生长发育;高CO₂浓度对小麦幼苗的生长发育有明显的促进作用,对分蘖数的影响更显著,干旱条件下高CO₂浓度使旱选3号和烟农19的分蘖数分别增加了61.0%和42.3%。两种水分条件下,高CO₂浓度显著增加了小麦幼苗的生物量,降低了幼苗叶片过氧化物酶和脯氨酸的含量。干旱条件下,高CO₂浓度表现出更好的“肥效作用”。此外,不同持绿型小麦品种对高CO₂浓度的响应存在差异,旱选3号对CO₂的响应更敏感。因此,在未来气候变化背景下,CO₂浓度升高时可以适当减少田间灌水量,合理利用水资源,还需注意选择适宜的品种。     

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.     

Molecular mapping of QTLs conferring stay-green in grain sorghum (Sorghum bicolor L. Moench).

Drought resistance is of enormous importance in crop production. The identification of genetic factors involved in plant response to drought stress provides a strong foundation for improving drought tolerance. Stay-green is a drought resistance trait in sorghum (Sorghum bicolor L. Moench) that gives plants resistance to premature senescence under severe soil moisture stress during the post-flowering stage. The objective of this study was to map quantitative trait loci (QTLs) that control the stay-green and chlorophyll content in sorghum. By using a restriction fragment length polymorphism (RFLP) map, developed from a recombinant inbred line (RIL) population, we identified four stay-green QTLs, located on three linkage groups. The QTLs (Stg1 and Stg2) are on linkage group A, with the other two, Stg3 and Stg4, on linkage groups D and J, respectively. Two stay-green QTLs, Stg1 and Stg2, explaining 13-20% and 20-30% of the phenotypic variability, respectively, were consistently identified in all trials at different locations in two years. Three QTLs for chlorophyll content (Chl1, Chl2, and Chl3), explaining 25-30% of the phenotypic variability were also identified under post-flowering drought stress. All coincided with the three stay-green QTL regions (Stg1, Stg2, and Stg3) accounting for 46% of the phenotypic variation. The Stg1 and Stg2 regions also contain the genes for key photosynthetic enzymes, heat shock proteins, and an abscisic acid (ABA) responsive gene. Such spatial arrangement shows that linkage group A is important for drought- and heat-stress tolerance and yield production in sorghum. High-resolution mapping and cloning of the consistent stay-green QTLs may help to develop drought-resistant hybrids and to understand the mechanism of drought-induced senescence in plants.