Genetic basis of drought resistance at reproductive stage in rice: separation of drought tolerance from drought avoidance

Drought tolerance (DT) and drought avoidance (DA) are two major mechanisms in drought resistance of higher plants. In this study, the genetic bases of DT and DA at reproductive stage in rice were analyzed using a recombinant inbred line population from a cross between an indica lowland and a tropical japonica upland cultivar. The plants were grown individually in PVC pipes and two cycles of drought stress were applied to individual plants with unstressed plants as the control. A total of 21 traits measuring fitness, yield, and the root system were investigated. Little correlation of relative yield traits with potential yield, plant size, and root traits was detected, suggesting that DT and DA were well separated in the experiment. A genetic linkage map consisting of 245 SSR markers was constructed for mapping QTL for these traits. A total of 27 QTL were resolved for 7 traits of relative performance of fitness and yield, 36 QTL for 5 root traits under control, and 38 for 7 root traits under drought stress conditions, suggesting the complexity of the genetic bases of both DT and DA. Only a small portion of QTL for fitness- and yield-related traits overlapped with QTL for root traits, indicating that DT and DA had distinct genetic bases.     

Quantitative trait loci for root-penetration ability and root thickness in rice: comparison of genetic backgrounds.

Drought is the major abiotic stress limiting rice (Oryza sativa) production and yield stability in rainfed lowland and upland ecosystems. Root systems play an important role in drought resistance. Incorporation of root selection criteria in drought resistance improvement is difficult due to lack of reliable and efficient screening techniques. Using a wax-petrolatum layer system simulated to compacted soil layers, root traits were evaluated in a doubled haploid (DH) population derived from the cross between 'IR64' and 'Azucena'. Twelve putative QTLs (quantitative trait loci) were detected by interval mapping comprising four QTLs for root-penetration ability, four QTLs for root thickness, two QTLs for penetrated root number, and two QTLs for total root number. These QTLs individually explained 8.4% to 16.4% of the phenotypic variation. No QTL was detected for maximum penetrated root length by interval mapping. One QTL located between RG104 and RG348 was found to influence both root-penetration ability and root thickness. QTLs for root-penetration ability and root thickness were compared across two populations, 'IR64'-'Azucena' and 'CO39'-'Moroberekan', and different testing conditions. The identified consistent QTLs could be used for marker-assisted selection for deep and thick roots with high root-penetration ability in rice.     

Mapping QTLs for Plant Phenology and Production Traits Using Indica Rice (Oryza sativa L.) Lines Adapted to Rainfed Environment

Drought is a major abiotic stress limiting rice production and yield stability in rainfed ecosystems. Identifying quantitative trait loci (QTL) for rice yield and yield components under water limited environments will help to develop drought resilient cultivars using marker assisted breeding (MAB) strategy. A total of 232 recombinant inbred lines of IR62266/Norungan were used to map QTLs for plant phenology and production traits under rainfed condition in target population of environments. A total of 79 QTLs for plant phenology and production traits with phenotypic variation ranging from 4.4 to 72.8% were detected under non-stress and drought stress conditions across two locations. Consistent QTLs for phenology and production traits were detected across experiments and water regimes. The QTL region, RM204-RM197-RM217 on chromosome 6 was linked to days to 50% flowering and grain yield per plant under both rainfed and irrigated conditions. The same genomic region, RM585-RM204-RM197 was also linked to harvest index under rainfed condition with positive alleles from Norungan, a local landrace. QTLs for plant production and drought resistance traits co-located near RM585-RM204-RM197-RM217 region on chromosome 6 in several rice genotypes. Thus with further fine mapping, this region may be useful as a candidate QTL for MAB, map-based cloning of genes and functional genomics studies for rainfed rice improvement.     

Molecular mapping of genomic regions harbouring QTLs for root and yield traits in sorghum (Sorghum bicolor L. Moench)

Root system is a vital part of plants for absorbing soil moisture and nutrients and it influences the drought tolerance. Identification of the genomic regions harbouring quantitative trait loci (QTLs) for root and yield traits, and the linked markers can facilitate sorghum improvement through marker-assisted selection (MAS) besides the deeper understanding of the plant response to drought stress. A population of 184 recombinant inbred lines (RILs), derived from E36-1 × SPV570, along with parents were phenotyped for component traits of yield in field and root traits in an above ground rhizotron. High estimates of heritability and genetic advance for all the root traits and for most of the yield traits, presents high scope for improvement of these traits by simple selection. A linkage map constructed with 104 marker loci comprising 50 EST-SSRs, 34 non-genic nuclear SSRs and 20 SNPs, and QTL analysis was performed using composite interval mapping (CIM) approach. A total of eight and 20 QTLs were mapped for root and yield related traits respectively. The QTLs for root volume, root fresh weight and root dry weight were found co-localized on SBI-04, supported by a positive correlation among these traits. Hence, these traits can be improved using the same linked markers. The lack of overlap between the QTLs of component traits of root and yield suggested that these two sets of parameters are independent in their influence and the possibility of combining these two traits might enhance productivity of sorghum under receding moisture condition.     

Locating genes associated with root morphology and drought avoidance in rice via linkage to molecular markers

This research was undertaken to identify and map quantitative trait loci (QTLs) associated with five parameters of rice root morphology and to determine if these QTLs are located in the same chromosomal regions as QTLs associated with drought avoidance/tolerance. Root thickness, root:shoot ratio, root dry weight per tiller, deep root dry weight per tiller, and maximum root length were measured in three replicated experiments (runs) of 203 recombinant inbred lines grown in a greenhouse. The lines were from a cross between indica cultivar Co39 andjaponica cultivar Moroberekan. The 203 RI lines were also grown in three replicated field experiments where they were drought-stressed at the seedling, early vegetative, and late-vegetative growth stage and assigned a visual rating based on leaf rolling as to their degree of drought avoidance/tolerance. The QTL analysis of greenhouse and field data was done using single-marker analysis (ANOVA) and interval analysis (Mapmaker QTL). Most QTLs that were identified were associated with root thickness, root/shoot ratio, and root dry weight per tiller, and only a few with deep root weight. None were reliably associated with maximum root depth due to genotype-by-experiment interaction. Root thickness and root dry weight per tiller were the characters found to be the least influenced by environmental differences between greenhouse runs. Correlations of root parameters measured in greenhouse experiments with field drought avoidance/tolerance were significant but not highly predictive. Twelve of the fourteen chromosomal regions containing putative QTLs associated with field drought avoidance/tolerance also contained QTLs associated with root morphology. Thus, selecting for Moroberekan alleles at marker loci associated with the putative root QTLs identified in this study may be an effective strategy for altering the root phenotype of rice towards that commonly associated with drought-resistant cultivars.     

Identification of QTL for increased fibrous roots in soybean

Drought stress adversely affects soybean at various developmental stages, which collectively results in yield reduction. Unpredictable rainfall has been reported to contribute about 36% to variation of yield difference between the rain-fed and irrigated fields. Among the drought resistance mechanisms, drought avoidance in genotypes with fibrous roots was recognized to be associated with drought resistance in soybean. Plant introduction PI416937 was shown to possess fibrous roots and has been used as a parent in breeding programs to improve soybean productivity. Little information is available on relative contribution and chromosomal location of quantitative trait loci (QTL) conditioning fibrous roots in soybean. To identify the genomic locations and genetic bases of this trait, a recombinant inbred line population was derived from a cross between PI416937 and ‘Benning’. To detect associated QTLs, phenotypic data were collected and analyzed for 2 years under rain-fed field conditions. The selective genotyping approach was used to reduce the costs and work associated with conducting the QTL analysis. A total of five QTLs were identified on chromosomes Gm01 (Satt383), Gm03 (Satt339), Gm04 (Sct_191), Gm08 (Satt429), and Gm20 (Sat_299), and together explained 51% of the variation in root score. Detected QTLs were co-localized with QTLs related to root morphology, suggesting that fibrous roots QTL may be associated with other morpho-physiological traits and seed yield in soybean. Genetic dissection of the fibrous roots trait at the individual marker loci will allow for marker-assisted selection to develop soybean genotypes with enhanced levels of fibrous roots.     

Marker-assisted selection to introgress rice QTLs controlling root traits into an Indian upland rice variety

A marker-assisted back-crossing (MABC) breeding programme was conducted to improve the root morphological traits, and thereby drought tolerance, of the Indian upland rice variety, Kalinga III. This variety, the recurrent parent in the MABC, had not previously been used for quantitative trait locus (QTL) mapping. The donor parent was Azucena, an upland japonica variety from Philippines. Five segments on different chromosomes were targeted for introgression; four segments carried QTLs for improved root morphological traits (root length and thickness) and the fifth carried a recessive QTL for aroma. Some selection was made at non-target regions for recurrent parent alleles. We describe the selection made in three backcross (BC) generations and two further crosses between BC₃ lines to pyramid (stack) all five target segments. Pyramids with four root QTLs were obtained in eight generations, completed in 6 years using 3,000 marker assays in a total of 323 lines. Twenty-two near-isogenic lines (NILs) were evaluated for root traits in five field experiments in Bangalore, India. The target segment on chromosome 9 (RM242-RM201) significantly increased root length under both irrigated and drought stress treatments, confirming that this root length QTL from Azucena functions in a novel genetic background. No significant effects on root length were found at the other four targets. Azucena alleles at the locus RM248 (below the target root QTL on chromosome 7) delayed flowering. Selection for the recurrent parent allele at this locus produced early-flowering NILs that were suited for upland environments in eastern India.     

Mapping QTLs associated with drought avoidance in upland rice

The identification of molecular markers linked to genes controlling drought resistance factors in rice is a necessary step to improve breeding efficiency for this complex trait. QTLs controlling drought avoidance mechanisms were analyzed in a doubled-haploid population of rice. Three trials with different drought stress intensities were carried out in two sites. Leaf rolling, leaf drying, relative water content of leaves and relative growth rate under water stress were measured on 105 doubled haploid lines in two trials and on a sub-sample of 85 lines in the third one. Using composite interval mapping with a LOD threshold of 2.5, the total number of QTLs detected in all trials combined was 11 for leaf rolling, 10 for leaf drying, 11 for relative water content and 10 for relative growth rate under stress. Some of these QTLs were common across traits. Among the eleven possible QTLs for leaf rolling, three QTLs (on chromosomes 1, 5 and 9) were common across the three trials and four additional QTLs (on chromosomes 3, 4 and 9) were common across two trials. One QTL on chromosome 4 for leaf drying and one QTL on chromosome 1 for relative water content were common across two trials while no common QTL was identified for relative growth rate under stress. Some of the QTLs detected for leaf rolling, leaf drying and relative water content mapped in the same places as QTLs controlling root morphology, which were identified in a previous study involving the same population. Some QTL identified here were also located similarly with other QTLs for leaf rolling as reported from other populations. This study may help to chose the best segments for introgression into rice varieties and improvement of their drought resistance.     

Mapping QTLs for root traits in a recombinant inbred population from two indica ecotypes in rice

Evaluation of root traits in rainfed lowland rice is very difficult. Molecular genetic markers could be used as an alternative strategy to phenotypic selection for the improvement of rice root traits. This research was undertaken to map QTLs associated with five root traits using RFLP and AFLP markers. Recombinant inbred lines (RILs) were developed from two indica parents, IR58821–23-B-1–2-1 and IR52561-UBN-1–1-2, that were adapted to rainfed lowland production systems. Using wax-petrolatum layers to simulate a hardpan in the soil, 166 RILs were evaluated for total root number (TRN), penetrated root number (PRN), root penetration index (RPI, the ratio of PRN to TRN), penetrated root thickness (PRT) and penetrated root length (PRL) under greenhouse conditions during the summer and the fall of 1997. A genetic linkage map of 2022 cM length was constructed comprising 303 AFLP and 96 RFLP markers with an average marker space of 5.0 cM. QTL analysis via interval mapping detected 28 QTLs for these five root traits, which were located on chromosomes 1, 2, 3, 4, 6, 7, 10 and 11. Individual QTLs accounted for between 6 and 27% of the phenotypic variation. Most of the favorable alleles were derived from the parent IR58821–23-B-1–2-1, which was phenotypically superior in root traits related to drought resistance. Three out of six QTLs for RPI were detected in both summer and fall experiments and they also were associated with PRN in both experiments. Out of eight QTLs for RPT, five were common in both seasons. Two genomic regions on chromosome 2 were associated with three root traits (PRN, PRT and RPI), whereas three genomic regions on chromosomes 2 and 3 were associated with two root traits (PRT and RPI). Two QTLs affecting RPI and two QTLs affecting PRT were also found in similar genomic regions in other rice populations. The consistent QTLs across genetic backgrounds and the common QTLs detected in both experiments should be good candidates for marker-assisted selection toward the incorporation of root traits in a drought resistance breeding program, especially for rainfed lowland rice. Received: 17 November 1999 / Accepted: 19 March 2000     

Mapping QTLs associated with drought avoidance in upland rice grown in the Philippines and West Africa

Localizing genes that contribute to drought avoidance in a quantitative way should enable the exploitation of these genes in breeding through marker-assisted selection, and may lead to the discovery of gene identity and function. Between 110 and 176 F₆ recombinant inbred lines from a mapping population derived from a cross of upland rice varieties Bala and Azucena have been evaluated for indicators of drought avoidance in sites in the Philippines and West Africa over two dry seasons. A molecular map with 102 RFLP, 34 AFLP and six microsatellite markers has been used to map (by composite interval mapping) quantitative trait loci (QTLs) for the visual scores of leaf rolling and leaf drying and leaf relative water content. QTLs were mapped for each site and across sites. A total of 17 regions were identified which contained QTLs with a LOD score greater than 3.2. For leaf rolling, Bala was the parent contributing the majority of positive alleles whilst for the other traits, Bala and Azucena contributed more evenly. Six of the 17 regions influenced more than one trait, explaining the phenotypic correlations between traits that were observed. Three QTLs appeared to be specific to the Philippines experiments. One QTL had opposing effects in the Philippines and West Africa. QTLs for relative water content were detected on chromosome 8, congruent with an osmotic adjustment QTL identified in another population. Only three of the QTLs identified here have not been reliably identified in the two other populations that have been screened for drought avoidance. By using several populations assessed for drought avoidance in different sites, the distribution and utility of QTLs for drought avoidance in rice is being elucidated.     

Toward understanding genetic mechanisms of complex traits in rice

Rice is the primary carbohydrate staple cereal feeding the world population. Many genes, known as quantitative trait loci (QTLs), con-trol most of the agronomically important traits in rice. The identification of QTLs controlling agricultural traits is vital to increase yield and meet the needs of the increasing human population, but the progress met with challenges due to complex QTL inheritance. To date,many QTLs have been detected in rice, including those responsible for yield and grain quality; salt, drought and submergence tolerance;disease and insect resistance; and nutrient utilization efficiency. Map-based cloning techniques have enabled scientists to successfully fine map and clone approximately seventeen QTLs for several traits. Additional in-depth functional analyses and characterizations of these genes will provide valuable assistance in rice molecular breeding.     

Quantitative trait loci associated with traits determining grain and stover yield in pearl millet under terminal drought-stress conditions

Drought stress during the reproductive stage is one of the most important environmental factors reducing the grain yield and yield stability of pearl millet. A QTL mapping approach has been used in this study to understand the genetic and physiological basis of drought tolerance in pearl millet and to provide a more-targeted approach to improving the drought tolerance and yield of this crop in water-limited environments. The aim was to identify specific genomic regions associated with the enhanced tolerance of pearl millet to drought stress during the flowering and grain-filling stages. Testcrosses of a set of mapping-population progenies, derived from a cross of two inbred pollinators that differed in their response to drought, were evaluated in a range of managed terminal drought-stress environments. A number of genomic regions were associated with drought tolerance in terms of both grain yield and its components. For example, a QTL associated with grain yield per se and for the drought tolerance of grain yield mapped on linkage group 2 and explained up to 23% of the phenotypic variation. Some of these QTLs were common across stress environments whereas others were specific to only a particular stress environment. All the QTLs that contributed to increased drought tolerance did so either through better than average maintenance (compared to non-stress environments) of harvest index, or harvest index and biomass productivity. It is concluded that there is considerable potential for marker-assisted backcross transfer of selected QTLs to the elite parent of the mapping population and for their general use in the improvement of pearl millet productivity in water-limited environments. Received: 15 November 2000 / Accepted: 12 April 2001     

Mapping QTLs of root morphological traits at different growth stages in rice

Roots are a vital organ for absorbing soil moisture and nutrients and influence drought resistance. The identification of quantitative trait loci (QTLs) with molecular markers may allow the estimation of parameters of genetic architecture and improve root traits by molecular marker-assisted selection (MAS). A mapping population of 120 recombinant inbred lines (RILs) derived from a cross between japonica upland rice 'IRAT109' and paddy rice 'Yuefu' was used for mapping QTLs of developmental root traits. All plant material was grown in PVC-pipe. Basal root thickness (BRT), root number (RN), maximum root length (MRL), root fresh weight (RFW), root dry weight (RDW) and root volume (RV) were phenotyped at the seedling (I), tillering (II), heading (III), grain filling (IV) and mature (V) stages, respectively. Phenotypic correlations showed that BRT was positively correlated to MRL at the majority of stages, but not correlated with RN. MRL was not correlated to RN except at the seedling stage. BRT, MRL and RN were positively correlated to RFW, RDW and RV at all growth stages. QTL analysis was performed using QTLMapper 1.6 to partition the genetic components into additive-effect QTLs, epistatic QTLs and QTL-by-year interactions (Q x E) effect. The results indicated that the additive effects played a major role for BRT, RN and MRL, while for RFW, RDW and RV the epistatic effects showed an important action and Q x E effect also played important roles in controlling root traits. A total of 84 additive-effect QTLs and 86 pairs of epistatic QTLs were detected for the six root traits at five stages. Only 12 additive QTLs were expressed in at least two stages. This indicated that the majority of QTLs were developmental stage specific. Two main effect QTLs, brt9a and brt9b, were detected at the heading stage and explained 19% and 10% of the total phenotypic variation in BRT without any influence from the environment. These QTLs can be used in breeding programs for improving root traits.     

Detection of Quantitative Trait Loci for Yield and Drought Tolerance Traits in Soybean Using a Recombinant Inbred Line Population

To investigate the genetic basis of drought tolerance in soybean ( Glycine max L. Merr.) a recombinant inbred population with 184 F_2:7:11 lines developed from a cross between Kefeng1 (drought tolerant) and Nannong1138-2 (drought sensitive) were tested under water-stressed and well-watered conditions in field and greenhouse trials. Traits measured included leaf wilting coefficient, excised leaf water loss and relative water content as indicators of plant water status and seed yield. A total of 40 quantitative trait loci (QTLs) were identified: 17 for leaf water status traits under drought stress and 23 for seed yield under well-watered and drought-stressed conditions in both field and greenhouse trials. Two seed yield QTLs were detected under both well-watered and drought-stressed conditions in the field on molecular linkage group H and D1b, while two seed yield QTLs on molecular linkage group C2 were found under greenhouse conditions. Several QTLs for traits associated with plant water status were identified in both field and greenhouse trials, including two leaf wilting coefficient QTLs on molecular linkage group A2 and one excised leaf water loss QTL on molecular linkage group H. Phenotypic correlations of traits suggested several QTLs had pleiotropic or location-linked associations. These results will help to elucidate the genetic basis of drought tolerance in soybean, and could be incorporated into a marker-assisted selection breeding program to develop high-yielding soybean cultivars with improved tolerance to drought stress.     

Dynamic QTL analysis of the Na+ content,K+ content,and Na+/K+ ratio in rice roots during the field growth under salt stress

Rice (Oryza sativa L.) is seriously impacted by global soil salinization. To determine the quantitative trait loci (QTLs) related to salt tolerance in rice roots, F_2:3 and BC₁F_2:3 populations derived from a cross between the cv. Dongnong 425 of high quality and yield and the salt-tolerant cv. Changbai 10, were studied at different development stages. Two genetic linkage maps of F_2:3 and BC1F_2:3 populations were constructed. A 66 mM NaCl solution was used to irrigate the field and to analyze the dynamic QTL of some rice root traits. Using unconditional and conditional QTL mapping methods, 30 unconditional QTLs and 16 conditional QTLs related to the 6 root traits were detected on the 9 rice chromosomes during different developmental stages. Fourteen pairs of unconditional and conditional QTLs were detected at the identical developmental stage in the identical population. A number of QTLs were detected at different developmental stages, however, many did not appear at the last stage. Remarkably, qRKC1 appeared continuously at multiple stages in both the populations suggesting its key role in regulating the salt tolerance of rice roots.     

Genetic dissection of root growth in rice (Oryza sativa L.). II: mapping quantitative trait loci using molecular markers

Drought is a major abiotic stress of upland rice, and good root growth has been associated with drought avoidance. We report on the genetic mapping of root growth traits in an F₂ population derived from two drought-resistant rice varieties, ‘Bala’ and ‘Azucena’. Restriction fragment length polymorphism (RFLP) between the parents was 32%, and a molecular map with 71 marker loci and 17 linkage groups covering 1280 cM was produced. Quantitative trait loci (QTLs) for eight root growth characteristics were mapped using phenotype data obtained in a hydroponic screen previously described in a companion paper. Using a significance threshold of LOD 2.4, we observed one QTL for maximum root length after 28 days growth on chromosome 11. It had a LOD score of 6.9, explained nearly 30% of the variation and appeared to be largely additive in effect. QTLs for maximum root length after 3, 7, 14 and 21 days of growth were also revealed. Some root-length QTLs, including that on chromosome 11, varied greatly with developmental stage. One QTL for root volume and two QTLs for adventitious root thickness were detected. No QTLs were detected for the length of cells in the mature (fully expanded) zone of adventitious root tips. The results obtained are discussed in the context of previous reports on mapping root growth parameters in rice.     

Studying the genetic basis of drought tolerance in sorghum by managed stress trials and adjustments for phenological and plant height differences

Managed environments in the form of well watered and water stressed trials were performed to study the genetic basis of grain yield and stay green in sorghum with the objective of validating previously detected QTL. As variations in phenology and plant height may influence QTL detection for the target traits, QTL for flowering time and plant height were introduced as cofactors in QTL analyses for yield and stay green. All but one of the flowering time QTL were detected near yield and stay green QTL. Similar co-localization was observed for two plant height QTL. QTL analysis for yield, using flowering time/plant height cofactors, led to yield QTL on chromosomes 2, 3, 6, 8 and 10. For stay green, QTL on chromosomes 3, 4, 8 and 10 were not related to differences in flowering time/plant height. The physical positions for markers in QTL regions projected on the sorghum genome suggest that the previously detected plant height QTL, Sb-HT9-1, and Dw2, in addition to the maturity gene, Ma5, had a major confounding impact on the expression of yield and stay green QTL. Co-localization between an apparently novel stay green QTL and a yield QTL on chromosome 3 suggests there is potential for indirect selection based on stay green to improve drought tolerance in sorghum. Our QTL study was carried out with a moderately sized population and spanned a limited geographic range, but still the results strongly emphasize the necessity of corrections for phenology in QTL mapping for drought tolerance traits in sorghum.     

QTL mapping for yield and photosynthetic related traits under different water regimes in wheat

The improvement for drought tolerance requires understanding of the genetic control of wheat (Triticum aestivum L.) reaction to drought. In this study, a set of 131 recombinant inbred lines of wheat were investigated under well-watered (WW) and drought stress (DS) environments across 2 years to map quantitative trait loci (QTLs) for yield and physiological traits. A total of 225 QTLs were detected, including 32 non-environment-specific loci that were significant in both DS and WW, one drought-specific locus and two watering-specific loci. Three consistently-expressed QTLs (QTkw-3A.2, QTss-1A, and QScn-7A.1) were identified in at least three environments and the QTkw-1D.1 was significant in DS across the 2 years. By unconditional and conditional QTL analysis, spike number per plant and kernel number per spike were more important than thousand-kernel weight for grain yield (GY) at the given genetic background. Meta-analysis identified 67 meta-QTLs that contained QTLs for at least two traits. High frequency co-location of QTLs was found among either the spike-related traits or the six physiological traits. Four photosynthesis traits (CHL, LWUE, P _N, and C _i) were co-located with GY and/or yield components on various MQTLs. The results provided QTLs that warrant further study for drought tolerance breeding and are helpful for understanding the genetic basis of drought tolerance and the genetic contribution of yield components to GY at individual QTL level in wheat.     

Comparison of QTLs for rice seedling morphology under different water supply conditions

The variation of seedling characteristics under different water supply conditions is strongly associated with drought resistance in rice (Oryza sativa L.) and a better elucidation of its genetics is helpful for improving rice drought resistance. Ninety-six doubled-haploid (DH)rice lines of an indica and japonica cross were grown in both flooding and upland conditions and QTLs for morphological traits at seedling stage were examined using 208 restriction fragment length polymorphism (RFLP) and 76 microsatellite (SSR) markers. A total of 32 putative QTLs were associated with the four seedling traits: average of three adventitious root lengths (ARL), shoot height (SH), shoot biomass (SW), and root to shoot dry weight ratio (RSR). Five QTLs detected were the same under control and upland conditions. The ratio between the mean value of the seedling trait under upland and flooding conditions was used for assessing drought tolerance. A total of six QTLs for drought tolerance were detected. Comparative analysis was performed for the QTLs detected in this case and those reported from two other populations with the same upland rice variety Azucena as parent. Several identical QTLs for seedling elongation across the three populations with the positive alleles from the upland rice Azucena were detected, which suggests that the alleles of Azucena might be involved in water stress-accelerated elongation of rice under different genetic backgrounds. Five cell wall-related candidate genes for OsEXP1, OsEXP2, OsEXP4, EXT, and EGase were mapped on the intervals carrying the QTLs for seedling traits.     

Linking drought-resistance mechanisms to drought avoidance in upland rice using a QTL approach: progress and new opportunities to integrate stomatal and mesophyll responses

The advent of saturated molecular maps promised rapid progress towards the improvement of crops for genetically complex traits like drought resistance via analysis of quantitative trait loci (QTL). Progress with the identification of QTLs for drought resistance-related traits in rice is summarized here with the emphasis on a mapping population of a cross between drought-resistant varieties Azucena and Bala. Data which have used root morphological traits and indicators of drought avoidance in field-grown plants are reviewed, highlighting problems and uncertainties with the QTL approach. The contribution of root-growth QTLs to drought avoidance appears small in the experiments so far conducted, and the limitations of screening methodologies and the involvement of shoot-related mechanisms of drought resistance are studied. When compared to Azucena, Bala has been observed to have highly sensitive stomata, does not roll its leaves readily, has a greater ability to adjust osmotically, slows growth more rapidly when droughted and has a lower water-use efficiency. It is also a semi-dwarf variety and hence has a different canopy structure. There is a need to clarify the contribution of the shoot to drought resistance from the level of the biochemistry of photosynthesis through stomatal behaviour and leaf anatomy to canopy architecture. Recent advances in studying the physical and biochemical processes related to water use and drought stress offer the opportunity to advance a more holistic understanding of drought resistance. These include the potential use of infrared thermal imaging to study energy balance, integrated and online stable isotope analysis to dissect processes involved in carbon dioxide fixation and water evaporation, and leaf fluorescence to monitor photosynthesis and photochemical quenching. Justification and a strategy for this integrated approach is described, which has relevance to the study of drought resistance in most crops.