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.     

Defining selection criteria to improve yield under drought

The many selection criteria that have been proposed to increase drought resistance of our crops have had little, if any, impact on improving crop yields in dry environments. There are several likely reasons for this lack of success. Some of these are: (i) criteria proposed have been related more to survival mechanisms under drought than to productivity, (ii) criteria are inappropriate to the target environment, and (iii) criteria are temporal and are therefore likely to have minimal impact on growth and yield over the entire lifecycle. Another important reason is that breeders have not been convinced the proposed criteria will be successful as they are too difficult to measure. On the other hand, empirical breeding programmes to improve yield under drought have been successful. Surprisingly, some of the greatest successes have been achieved by breeding in environments where water is non-limiting. This paper reviews breeding approaches to improve yield under drought. It focuses on critical factors that must be considered to identify likely plant attributes that can be targeted. These factors, their link with yield, the nature of the target environment, the level of organisation where the trait is expressed are discussed. Three quite different examples are given to emphasize the above considerations and which show substantial promise in targeting traits to improve yield under drought. They are drought at flowering, improving transpiration efficiency and improving early leaf area development.     

Genetics of yield component traits under salt stress at flowering stage and selection of salt tolerant pre-breeding lines for rice improvement

Genetica - Rice is highly vulnerable to salt stress at both seedling and flowering stage. While research efforts largely focused on seedling stage salinity tolerance, flowering stage salt tolerance...     

Effect of soil mechanical impedance on root growth of two rice varieties under field drought stress

Two upland rice varieties, Azucena and Bala, were screened for root growth under droughted and irrigated treatments in two field sites at the West Africa Rice Development Association (WARDA) experimental farm, Côte d’Ivoire, during the dry season of 1999/2000. The sites were chosen to represent contrasting soil profile penetration resistance (PR) characteristics on upland sites, although both were relatively impeding. The number of nodal root axes per unit area passing through horizontal transects (root density) was counted at 35, 56, 77 and 98 days after sowing (DAS) at 10 cm depth intervals. Azucena consistently maintained a greater root density than Bala and a greater proportion of Azucena roots grew to 30 cm depth (22.7% vs. 8.4% at 77 DAS). There was little detectable effect of water regime on root distribution but evidence of lower root numbers at depths below 20 cm in the higher PR site was revealed. A site by variety by soil depth interaction suggests that Azucena roots are more strongly affected by very high PR than those of Bala. PR between 0–30 cm depth increased greatly with decreasing soil water content during the drought as the soil dried. This increase is likely to have prevented or greatly impaired further nodal root growth within this layer. At 40 cm depth, PR was high (3–4 MPa) but did not increase during the drought. At this depth root growth rate was likely to be greatly reduced despite the availability of water. These results demonstrate that varietal differences in root morphology characterised in the laboratory can be also detected in impeding field soils as differences in the density of roots at depth. Relatively poor root growth in these fields in the absence of drought was probably due to the high mechanical impedance and/or the physiological stress of the plants in the dry season. Our results indicate that high mechanical impedance was a more fundamental constraint on root growth than soil water availability during the drought. Thus, varietal differences in root penetration ability might be very important for drought avoidance in soils of this type.     

Meta-analysis of quantitative trait loci for grain yield and component traits under reproductive-stage drought stress in an upland rice population

A recombinant inbred population developed from a cross between high-yielding lowland rice (Oryza sativa L.) subspecies indica cv. IR64 and upland tropical rice subspecies japonica cv. Cabacu was used to identify quantitative trait loci (QTLs) for grain yield (GY) and component traits under reproductive-stage drought stress. One hundred fifty-four lines were grown in field trials in Indonesia under aerobic conditions by giving surface irrigation to field capacity every 4 days. Water stress was imposed for a period of 15 days during pre-flowering by withholding irrigation at 65 days after seeding. Leaf rolling was scored at the end of the stress period and eight agronomic traits were evaluated after recovery. The population was also evaluated for root pulling force, and a total of 201 single nucleotide polymorphism markers were used to construct the molecular genetic linkage map and QTL mapping. A QTL for GY under drought stress was identified in a region close to the sd1 locus on chromosome 1. QTL meta-analysis across diverse populations showed that this QTL was conserved across genetic backgrounds and co-localized with QTLs for leaf rolling and osmotic adjustment (OA). A QTL for percent seed set and grains per panicle under drought stress was identified on chromosome 8 in the same region as a QTL for OA previously identified in three different populations.     

Fine mapping of QTLs for rice grain yield under drought reveals sub-QTLs conferring a response to variable drought severities

Fine-mapping studies on four QTLs, qDTY(2.1), qDTY(2.2), qDTY(9.1) and qDTY(12.1), for grain yield (GY) under drought were conducted using four different backcross-derived populations screened in 16 experiments from 2006 to 2010. Composite and bayesian interval mapping analyses resolved the originally identified qDTY(2.1) region of 42.3 cM into a segment of 1.6 cM, the qDTY(2.2) region of 31.0 cM into a segment of 6.7 cM, the qDTY(9.1) region of 32.1 cM into two segments of 9.4 and 2.4 cM and the qDTY(12.1) region of 10.6 cM into two segments of 3.1 and 0.4 cM. Two of the four QTLs (qDTY(9.1) and qDTY(12.1)) having effects under varying degrees of stress severity showed the presence of more than one region within the original QTL. The study found the presence of a donor allele at RM262 within qDTY(2.1) and RM24334 within qDTY(9.1) showing a negative effect on GY under drought, indicating the necessity of precise fine mapping of QTL regions before using them in marker-assisted selection (MAS). However, the presence of sub-QTLs together in close vicinity to each other provides a unique opportunity to breeders to introgress such regions together as a unit into high-yielding drought-susceptible varieties through MAS.     

Morphophysiology, morphoanatomy, and grain yield under field conditions for two maize hybrids with contrasting response to drought stress

In the northern region of the state of Minas Gerais, lack of rainfall limits crop production in the field, which is possible only with irrigation. Agricultural and physiological practices have been intensively searched to overcome drought effects and consequently increase production. In this context, the objective of this study was to characterize morphophysiological and morphoanatomical changes and evaluate the attributes of grain yield under field conditions in two hybrids contrasting for drought tolerance. The experiment was carried out for 2 years (2010 and 2011) and the water deficit was imposed by stopping irrigation for 22 days at the pre-flowering stage. At the end of the stress treatment, leaf and root anatomy and morphophysiological characteristics (leaf water potential, chlorophyll content, percentage of dry leaves, leaf area, stomatal conductance, chlorophyll fluorescence, and anthesis-silking interval) were evaluated. For a better interpretation of tolerance of the hybrids in the evaluated characteristics, an index was used stress index. Hybrid DKB 390 (tolerant) surpassed hybrid BRS 1030 (sensitive) in grain yield. Furthermore, it presented lower percentage of dry leaves, higher flowering synchronization, higher stomatal conductance, and higher Fv/Fm relationship. In the root, DKB 390 showed higher amount of aerenchyma in the cortex, an increase of exodermis width, and numerous metaxylem with smaller diameter. In the leaf, it presented higher number of stomata and smaller distance between the vascular bundles in the leaf blade. The study concluded that significant morphophysiological and morphoanatomical changes, which are related to drought tolerance, occurred in DKB 390, leading to a higher yield in the field.     

Transmission of important chromosomal regions under selection revealed in rice pedigree breeding programs

Genetic analysis across a whole plant genome based on pedigree information offers considerable potential for enhancing genetic gain from plant breeding programs through quantitative trait loci (QTL) mapping and marker-assisted selection. Here, we report its application for graphically genotyping varieties used in Chinese japonica rice (Oryza sativa L.) pedigree breeding programs. We identified 34 important chromosomal regions from the founder parent that are under selection in the breeding programs, and by comparing donor genomic regions that are under selection with QTL locations of agronomic traits, we found that QTL clustered in important genomic regions, in accordance with association analyses of natural populations and other previous studies. The convergence of genomic regions under selection with QTL locations suggests that donor genomic regions harboring key genes/QTL for important agronomic traits have been selected by plant breeders since the 1950s from the founder rice plants. The results provide better understanding of the effects of selection in breeding programs on the traits of rice cultivars. They also provide potentially valuable information for enhancing rice breeding programs through screening candidate parents for targeted molecular markers, improving crop yield potential and identifying suitable genetic material for use in future breeding programs.     

QTLs for cell-membrane stability mapped in rice (Oryza sativa L.) under drought stress

Cell-membrane stability (CMS) is considered to be one of the major selection indices of drought tolerance in cereals. In order to determine which genomic region is responsible for CMS, 104 rice (Oryza sativa L.) doubled haploid (DH) lines derived from a cross between CT9993–5-10–1-M and IR62266-42–6-2 were studied in the greenhouse in a slowly developed drought stress environment. Drought stress was induced on 50-day-old plants by withholding water. The intensity of stress was assessed daily by visual scoring of leaf wilting and by measuring leaf relative water content (RWC). The leaf samples were collected from both control (well-watered) and stressed plants (at 60–65% of RWC), and the standard test for CMS was carried out in the laboratory. There was no significant difference (P>0.05) in RWC between the two parental lines as well as among the 104 lines, indicating that all the plants were sampled at a uniform stress level. However, a significant difference (P<0.05) in CMS was observed between the two parental lines and among the population. No significant correlation was found between CMS and RWC, indicating that the variation in CMS was genotypic in nature. The continuous distribution of CMS and its broad-sense heritability (34%) indicates that CMS should be polygenic in nature. A linkage map of this population comprising of 145 RFLPs, 153 AFLPs and 17 microsatellite markers was used for QTL analysis. Composite interval mapping identified nine putative QTLs for CMS located on chromosomes 1, 3, 7, 8, 9, 11 and 12. The amount of phenotypic variation that was explained by individual QTLs ranged from 13.4% to 42.1%. Four significant (P<0.05) pairs of digenic interactions between the detected QTLs for CMS were observed. The identification of QTLs for this important trait will be useful in breeding for the improvement of drought tolerance in rice. This is the first report of mapping QTLs associated with CMS under a natural water stress condition in any crop plants. Received: 8 September 1999 / Accepted: 13 October 1999     

Towards molecular breeding and improvement of rice in China

China is the largest producer and consumer of rice in the world and a pioneer in applying hybrid rice technology. Although hybrid rice has contributed greatly to Chinese agriculture in the past decades, its potential to improve grain quality further is being questioned. However, to meet the challenges posed by severe crop damage by pests and diseases, the extensive use of pesticides and chemical fertilizers, and a shortage of water and energy, more elite rice cultivars are needed. In recent years, China has seen continued improvements in rice genetics, powered by functional genomics as a way forward to safeguard its rice production. Here, we briefly review the current status of rice breeding in China through strategies integrating hybrid rice technology, molecular marker-assisted breeding, functional genomics and genetically modified rice.     

Seasonal radiation interception, dry matter production and yield determination for a semi-leafless pea (Pisum sativum) breeding selection under contrasting field conditions

Radiation interception, dry matter accumulation, flower and pod production and yield were measured for a semi-leafless pea (Pisum sativum) breeding selection (BS3) on three contrasting sites. Differences in soil moisture availability were largely responsible for a three-fold difference in yield between sites. Radiation interception was related to dry matter production by calculating photosynthetic efficiencies. In the absence of lodging, crop canopies converted intercepted radiation into dry matter with constant efficiency (?) throughout the season; under conditions of moisture stress ? was reduced. Serious lodging during the post-flowering period on one site resulted in a mean seasonal photosynthetic efficiency (?) 17% lower than ?. The ability of the pea crop canopy to intercept radiation was related also to yield components.     

Proteomic analysis of rice leaf sheath during drought stress

Drought is one of the most severe limitations on the productivity of rainfed lowland and upland rice. To investigate the initial response of rice to drought stress, changes in protein expression were analyzed using a proteomic approach. Two-week-old rice seedlings were exposed to drought conditions from 2 to 6 days, and proteins were extracted from leaf sheaths, separated by two-dimensional polyacrylamide gel electrophoresis and stained with Coomassie brilliant blue. After drought stress for 2 to 6 days, 10 proteins increased in abundance and the level of 2 proteins decreased. The functional categories of these proteins were identified as defense, energy, metabolism, cell structure, and signal transduction. In addition to drought stress, accumulations of protein were analyzed under several different stress conditions. The levels of an actin depolymerizing factor, a light harvesting complex chain II, a superoxidase dismutase and a salt-induced protein were changed by drought and osmotic stresses, but not cold or salt stresses, or abscisic acid treatment. The effect of drought stress on protein in the leaf sheaths of drought-tolerant rice cultivar was also analyzed. The light harvesting complex chain II and the actin depolymerizing factor were present at high levels in a drought-tolerant rice cultivar before stress application. With drought stress, actin depolymerizing factor was expressed in leaf blades, leaf sheaths, and roots. These results suggest that actin depolymerizing factor is one of the target proteins induced by drought stress.     

Exploitation of synthetic-derived wheats through osmotic stress responses for drought tolerance improvement

The development of drought tolerant wheat cultivars has been slow due to lack of understanding the diagnostic physiological parameters associated with improved productivity under water stress. We evaluated responses to PEG induced osmotic stress under hydroponics in D-genome synthetic derived and bread wheat germplasm with the main aim to unravel and identify some promising attributes having role in stress tolerances. Genotypes used in this study differed in their morpho-physiological and biochemical attributes. Tolerant genotypes exhibited the ability to ameliorate harmful effects of PEG induced osmotic stress through better osmotic adjustment achieved through substantial relative water content (RWC), lowered osmotic potential, relatively stable root length having maximum water extraction capacity, significant increase in osmoprotectant concentration and relatively enhanced antioxidant activities. The results clearly revealed the importance of synthetic derivatives over check cultivars and conventional wheats in terms of osmotic stress responses. Interestingly, synthetic-derived advanced lines with Aegilops tauschii in its parentage including AWL-02, AWL-04 and AWL-07 proved superior over the best rainfed check cultivar (Wa-01). It was concluded that synthetic-derived wheats has great potential to improve a range of stress adaptive traits. It could, therefore, be recommended to be a useful strategy for allowing modern bread wheat to become adapted to a wider range of environments in future climate change scenarios.     

Phytohormones and rice crop yield: strategies and opportunities for genetic improvement

To feed an estimated world population of 8.9 billion by 2050, strategies for increasing grain production must be developed. Several agronomically important traits for increasing yield, such as plant height, grain number, and leaf erectness, have recently been characterized in rice (Oryza sativa L.). These traits are regulated primarily by three phytohormones: gibberellins, cytokinins, and brassinosteroids. The control of biosynthesis and degradation of these key phytohormones is discussed in terms of its importance for normal plant growth. Genes involved in the biosynthesis and regulation of these phytohormones can be used to develop effective strategies to increase grain yield. Genetic manipulation of phytohormone-related gene expression is thus a practical strategy to generate high-yielding transgenic plants through the modification of levels and profile of endogenous phytohormones.