SummaryHybrid breeding is the main strategy for improving productivity in many crops, especially ... more SummaryHybrid breeding is the main strategy for improving productivity in many crops, especially in rice and maize. Genomic hybrid breeding is a technology that uses whole‐genome markers to predict future hybrids. Predicted superior hybrids are then field evaluated and released as new hybrid cultivars after their superior performances are confirmed. This will increase the opportunity of selecting true superior hybrids with minimum costs. Here, we used genomic best linear unbiased prediction to perform hybrid performance prediction using an existing rice population of 1495 hybrids. Replicated 10‐fold cross‐validations showed that the prediction abilities on ten agronomic traits ranged from 0.35 to 0.92. Using the 1495 rice hybrids as a training sample, we predicted six agronomic traits of 100 hybrids derived from half diallel crosses involving 21 parents that are different from the parents of the hybrids in the training sample. The prediction abilities were relatively high, varying f...
Hybrid rice varieties can outyield the best inbred varieties by 15 – 30% with appropriate managem... more Hybrid rice varieties can outyield the best inbred varieties by 15 – 30% with appropriate management. However, hybrid rice requires more inputs and management than inbred rice to realize a yield advantage in high-yielding environments. The development of stress-tolerant hybrid rice with lowered input requirements could increase hybrid rice yield relative to production costs. We used genomic prediction to evaluate the combining abilities of 564 stress-tolerant lines used to develop Green Super Rice with 13 male sterile lines of the International Rice Research Institute for yield-related traits. We also evaluated the performance of their F1 hybrids. We identified male sterile lines with good combining ability as well as F1 hybrids with potential further use in product development. For yield per plant, accuracies of genomic predictions of hybrid genetic values ranged from 0.490 to 0.822 in cross-validation if neither parent or up to both parents were included in the training set, and b...
The underground reserve (root) has been an uncharted research territory with its untapped genetic... more The underground reserve (root) has been an uncharted research territory with its untapped genetic variation yet to be exploited. Identifying ideal traits and breeding new rice varieties with efficient root system architecture (RSA) has great potential to increase resource-use efficiency and grain yield, especially under direct-seeded rice, by adapting to aerobic soil conditions. In this review, we tried to mine the available research information on the direct-seeded rice (DSR) root system to highlight the requirements of different root traits such as root architecture, length, number, density, thickness, diameter, and angle that play a pivotal role in determining the uptake of nutrients and moisture at different stages of plant growth. RSA also faces several stresses, due to excess or deficiency of moisture and nutrients, low or high temperature, or saline conditions. To counteract these hindrances, adaptation in response to stress becomes essential. Candidate genes such as early ro...
We studied variation in adaptive traits and genetic association to understand the low P responses... more We studied variation in adaptive traits and genetic association to understand the low P responses, including the symbiotic association of arbuscular mycorrhizal (AM) fungal colonization in Oryza species (O. sativa, O. nivara, and O. rufipogon). In the present experiment, we performed the phenotypic variability of the morphometric and geometric traits for P deficiency tolerance and conducted the association studies in GLM and MLM methods. A positive association between the geometric trait of the top-view area and root traits suggested the possibility of exploring a non-destructive approach in screening genotypes under low P. The AMOVA revealed a higher proportion of variation among the individuals as they belonged to different species of Oryza and the NM value was 2.0, indicating possible gene flow between populations. A sub-cluster with superior-performing accessions had a higher proportion of landraces (42.85%), and O. rufipogon (33.3%) was differentiated by four Pup1-specific mark...
Achieving sufficient food with finite resources and leaving lesser impact on the environment to f... more Achieving sufficient food with finite resources and leaving lesser impact on the environment to feed the growing global population has always been a great challenge. Here we review the concept and practices of Green Super Rice (GSR) that has led to a paradigm shift in the goals of crop genetic improvement and models of food production for promoting sustainable agriculture. The momentous achievements and global deliveries of GSR have been fueled through the integration of abundant genetic resources, functional gene discoveries, innovative breeding techniques with precise gene and whole-genome selections, and efficient agronomic management for promoting resource-saving and environment-friendly crop production systems. We also provide perspectives of new horizons in future genomic breeding technologies towards delivering green and nutritious crop varieties to further enhance the development of green agriculture to better nourish the world population.
Nitrogen is one of the most important nutrients for rice growth and development. Breeding of nitr... more Nitrogen is one of the most important nutrients for rice growth and development. Breeding of nitrogen deficiency tolerance (NDT) variety is considered to be the most economic measure to solve the constrain of low nitrogen stress on grain yield in rice. An interconnected breeding (IB) population of 497 lines developed using Huanghuazhan (HHZ) as the recurrent parent and eight elite lines as the donor parents were tested for five traits including grain yield, biomass, harvest index, thousand grain weight, and spikelet fertility under two nitrogen treatments in three growing seasons. Association analysis using 7,388 bins generated by sequencing identified a total of 14, 14, and 12 QTLs for the five traits under low nitrogen (LN), normal nitrogen (NN), and LN/NN conditions, respectively, across three seasons. Favorable alleles were dissected for the 40 QTLs at the 10 NDT regions, and OM1723 was considered as the most important parent with the highest frequency of favorable alleles contr...
Rice remains a major staple food source for the rapidly growing world population. However, regula... more Rice remains a major staple food source for the rapidly growing world population. However, regular occurrences of carcinogenic arsenic (As) minerals in waterlogged paddy topsoil pose a great threat to rice production and consumers across the globe. Although As contamination in rice has been well recognized over the past two decades, no suitable rice germplasm had been identified to exploit in adaptive breeding programs. Therefore, this current study identified suitable rice germplasm for As tolerance and exclusion based on a variety of traits and investigated the interlinkages of favorable traits during different growth stages. Fifty-three different genotypes were systematically evaluated for As tolerance and accumulation. A germination screening assay was carried out to identify the ability of individual germplasm to germinate under varying As stress. Seedling-stage screening was conducted in hydroponics under varying As stress to identify tolerant and excluder genotypes, and a fie...
BackgroundImproving performance of rice under drought stress has potential to significant impact ... more BackgroundImproving performance of rice under drought stress has potential to significant impact on rice productivity. Previously we reported that OsLG3 positively control rice grain length and yield.ResultsIn this study, we found that OsLG3 was more strongly expressed in upland rice compared to lowland rice under drought stress condition. Candidate gene association analysis showed that the natural variation in OsLG3 was associated with tolerance to water deficit stress in germinating rice seeds. Transgenic rice with enhanced OsLG3 expression exhibited improved tolerance to drought and that is most likely due to enhanced ROS scavenging efficiency. Phylogenetic analysis and pedigree records indicated that the tolerant allele of OsLG3 has potential to improve drought tolerance of japonica rice.ConclusionsCollectively, our work revealed that the natural variation of OsLG3 contributes to rice drought tolerance and the elite allele of OsLG3 is a promising genetic resource for the develop...
To develop superior rice varieties with improved yield in most rainfed areas of Asia/Africa, we s... more To develop superior rice varieties with improved yield in most rainfed areas of Asia/Africa, we started an introgression-breeding program for simultaneously improving yield and tolerances of multiple abiotic stresses. Using eight BC1 populations derived from a widely adaptable recipient and eight donors plus three rounds of phenotypic selection, we developed 496 introgression lines (ILs) with significantly higher yield under drought, salt and/or non-stress conditions in 5 years. Six new varieties were released in the Philippines and Pakistan and many more are being evaluated in multi-location yield trials for releasing in several countries. Marker-facilitated genetic characterization revealed three interesting aspects of the breeding procedure: (1) the donor introgression pattern in specific BC populations was characteristic; (2) introgression frequency in different genomic regions varied considerably, resulting primarily from strong selection for the target traits; and (3) signific...
Cold stress is one of the major abiotic stresses that impede rice production. A interconnected br... more Cold stress is one of the major abiotic stresses that impede rice production. A interconnected breeding (IB) population consisted of 497 advanced lines developed using HHZ as the recurrent parent and eight diverse elite indica lines as the donors were used to identify stably expressed QTLs for CT at the booting stage. A total of 41,754 high-quality SNPs were obtained through re-sequencing of the IB population. Phenotyping was conducted under field conditions in two years and three locations. Association analysis identified six QTLs for CT on the chromosomes 3, 4 and 12. QTL qCT-3-2 that showed stable CT across years and locations was fine-mapped to an approximately 192.9 kb region. Our results suggested that GWAS applied to an IB population allows better integration of gene discovery and breeding. QTLs can be mapped in high resolution and quickly utilized in breeding.
Interaction between rice (Oryza sativa L.) and its bacterial blight (BB) pathogen Xanthomonas ory... more Interaction between rice (Oryza sativa L.) and its bacterial blight (BB) pathogen Xanthomonas oryzae pv. oryzae (Xoo) is the model system between monocot plants and their bacterial pathogens. To understand the genome-wide interactions between Xoo and rice resulting from a broad-spectrum hypersensitive reaction (BSHR) mediated by a new rice resistance (R) gene, Xa39, comparative dynamic transcriptomic profiles in the incompatible and compatible interactions were investigated using three related rice lines and a highly virulent Xoo isolate, PXO99. Large numbers of rice and Xoo differentially expressed genes (DEGs) were identified in comparisons between the incompatible interactions and compatible ones, suggesting the gene network consisted of 27 genes in four groups of distinct functions involving leading to BSHR of rice in the sequential events from the avrXa39 ´ Xa39 interaction ® signal recognition and transduction ® protein modification ® programmed cell death. Correspondingly, several groups of Xoo genes expressed or upregulated specifically in the incompatible interaction were those for type III secretion system (T3SS) and type III secretion effectors (T3SEs). Combined evidence suggests LOC_Os11g37759, one of a two-member CC-NBS-LRR gene family on rice chromosome 11, as the most likely candidate for Xa39 in rice and two genes, XopN and XopX involved in T3SS of Xoo, as the most likely candidate genes for the corresponding avrXa39 in Xoo. Our transcriptome data and identified rice and Xoo DEGs provided a valuable source of information for future investigations on the Xoo-rice interactions. B acterial blight caused by Xanthomonas oryzae pv. oryzae (Xoo) is one of the most widely distributed rice diseases in tropical Asia and Africa, causing severe rice yield loses as most high-yielding rice cultivars are highly susceptible (NiÑo-Liu et al., 2006). Developing cultivars resistant to BB is the most economic and effective method to control the disease. Understanding of the molecular mechanisms underlying interactions between host and pathogen is required for successful knowledge-based rice improvement.
Background: Bacterial blight, caused by Xanthomonas oryzae pv. oryzae (Xoo), is a devastating ric... more Background: Bacterial blight, caused by Xanthomonas oryzae pv. oryzae (Xoo), is a devastating rice disease worldwide. Xa39 is a resistance (R) gene with a broad-spectrum hypersensitive response (BSHR) to Xoo. Nevertheless, the molecular mechanisms of resistance mediated by Xa39 remain unclear. In this study, the transcriptome profiling of a rice line carrying Xa39 and its parents at the early stage of Xoo infection were investigated. Results: A rice introgression line H471 carrying Xa39 exhibited a typical local hypersensitive response phenotype, accompanied by programmed cell death after inoculation with the Xoo Philippines' race 9b. Transcriptome profiling of H471 and its parents at 1 and 2 days post-inoculation was performed using RNA sequencing. In total, 306 differentially expressed genes (DEGs) were identified in H471 compared with its recurrent parent Huang-Hua-Zhan after inoculation with Xoo. Among them, 121 (39.5%) genes, with functional enrichments that were related to defense response, protein amino acid phosphorylation, and apoptosis, were found to be constitutively expressed. The other 185 (60.5%) genes, with GO terms that belonged to defense response, were significantly responsive to Xoo infection in H471. Ten up-regulated and 12 down-regulated genes encoding intracellular immune receptors were identified in H471 compared with Huang-Hua-Zhan. LOC_Os11g37759, which was located in the fine-mapping region harboring Xa39, is a Xa39 candidate gene. The putative BSHR-related co-regulatory networks were constructed using 33 DEGs from four functional groups, including gibberellic acid receptors and brassinosteroid regulators, which were differentially co-expressed with LOC_Os11g37759 in infected H471. Our results indicated that there might be cross-talk between the Xa39-mediated signal transduction cascades and the GA/BR signaling pathway, and that the defense mechanism was related to diverse kinases, transcription factors, post-translational regulation, and R genes. Conclusions: The present study provides the comprehensive transcriptome profile of a rice introgression line carrying Xa39 and its parents, and identifies a set of DEGs involved in BSHR mediated by Xa39. These data provide novel insights into the regulatory networks of plant disease resistance mediated by R genes, and the identified DEGs will serve as candidates for Xa39 cloning and for further understanding the molecular mechanism of BSHR.
ABSTRACT Four improved indica varieties adapted to high altitude regions namely K 39-2, lET 11988... more ABSTRACT Four improved indica varieties adapted to high altitude regions namely K 39-2, lET 11988, Himdhan and lET 11990 were treated with ethyl methane sulfonate, sodium azide and gamma rays with the objective of determining the most efficient mutagen and most responsive genotype for induction of mutations affecting predominantly reproductive parts and their functioning in rice. Frequency and spectrum of mutations affecting floral traits computed on Ml plant progeny and M2 population basis revealed the combination of the efficient sodium azide and the most responsive variety lET 11990 to be ideal for recovering highest number of selectively male sterile mutations. Among three kinds of male steriles viz., typical abortive, sporogenous and deformed male sex types recovered, critical study of abortive type helped identify several temperature sensitive genic male steriles, two of which are being improved for commercial exploitation in two-line hybrid breeding.
Proceedings of the National Academy of Sciences, 2006
Complete resistance (CR) and partial resistance (PR) of rice ( Oryza sativa L.) to its bacterial ... more Complete resistance (CR) and partial resistance (PR) of rice ( Oryza sativa L.) to its bacterial pathogen, Xanthomonas oryzae pv. oryzae ( Xoo ), was genetically dissected by using 2 mapping populations and 10 Xoo races. Two CR genes, 50 quantitative resistance loci, and 60 digenic interactions were identified, which showed various degrees of race specificity to the Xoo races. The complex epistasis between these loci led us to the discovery of complex genetic networks underlying the rice defensive system to Xoo . The networks consisted of two major components: one representing interactions between alleles at the R loci of rice and alleles at the corresponding avirulence loci of Xoo for CR and the other comprising interactions between quantitative resistance loci in rice and their corresponding aggressiveness loci in Xoo for PR. The race specificity of PR and its strong genetic overlap with CR indicate that PR is essentially “weaker” CR. The genetic networks discovered are expected t...
Great progress has been made in genetic dissection of quantitative trait variation during the pas... more Great progress has been made in genetic dissection of quantitative trait variation during the past two decades, but many studies still reveal only a small fraction of quantitative trait loci (QTLs), and epistasis remains elusive. We integrate contemporary knowledge of signal transduction pathways with principles of quantitative and population genetics to characterize genetic networks underlying complex traits, using a model founded upon one-way functional dependency of downstream genes on upstream regulators (the principle of hierarchy) and mutual functional dependency among related genes (functional genetic units, FGU). Both simulated and real data suggest that complementary epistasis contributes greatly to quantitative trait variation, and obscures the phenotypic effects of many 'downstream' loci in pathways. The mathematical relationships between the main effects and epistatic effects of genes acting at different levels of signaling pathways were established using the quantitative and population genetic parameters. Both loss of function and ''coadapted'' gene complexes formed by multiple alleles with differentiated functions (effects) are predicted to be frequent types of allelic diversity at loci that contribute to the genetic variation of complex traits in populations. Downstream FGUs appear to be more vulnerable to loss of function than their upstream regulators, but this vulnerability is apparently compensated by different FGUs of similar functions. Other predictions from the model may account for puzzling results regarding responses to selection, genotype by environment interaction, and the genetic basis of heterosis.
Three populations with a total of 125 BC2F3:4 introgression lines (ILs) selected for high yields ... more Three populations with a total of 125 BC2F3:4 introgression lines (ILs) selected for high yields from three BC2F2 populations were used for genetic dissection of rice yield and its related traits. The progeny testing in replicated phenotyping across two environments and genotyping with 140 polymorphic simple sequence repeat markers allowed the identification of 21 promising ILs that had significantly higher yields than the recurrent parent Shuhui527 (SH527). A total of 94 quantitative trait loci (QTL) were identified using the selective introgression method based on Chi-squared (χ2) and multi-locus probability tests and the RSTEP-LRT method based on stepwise regression. These QTL were mostly mapped to 12 clusters on seven rice chromosomes. Several important properties of the QTL affecting grain yield (GY) and its related traits were revealed. The first one was the presence of strong and frequent non-random associations between or among QTL that affect low-heritability traits (GY and spikelet number per panicle, SN) in the ILs with high trait values. Second, beneficial alleles at 88.9 % GY and 75 % SN QTL for increased productivity were from the donors, suggesting that direct phenotypic selection for high yield in our introgression breeding program was a powerful way to transfer beneficial alleles at many loci from the donors into SH527. Third, most QTL were in clusters with large effects on multiple traits, which should be the focal points in further investigations and marker-assisted selection in rice. The majority of the QTL identified were expressed only in one of the environments, suggesting that differential expression of QTL in different environments is the primary genetic basis of genotype × environment interaction. Finally, a large variation in both the direction and magnitude of QTL effects was detected for different donor alleles at seven QTL in the same genetic background and environments. This finding suggests the possible presence of functional diversity among the donor alleles at these loci. The promising ILs and QTL identified provide valuable materials and genetic information for further improving the yield potential of SH527, which is a backbone restorer of hybrid rice in China.
Drought is the most important factor limiting rice productivity in the rainfed areas of Asia. In ... more Drought is the most important factor limiting rice productivity in the rainfed areas of Asia. In this study, 48 pyramiding lines (PLs) and their recurrent parent, IR64, were evaluated over two years for their yield performances and related traits under severe drought stress at the reproductive stage (RS), the vegetative stage (VS) and irrigated control in order to understand the relationship between drought tolerance (DT) and yield potential (YP) in rice and their underlying mechanisms. When compared with IR64, all PLs had significantly improved DT to RS and 36 PLs also had significantly improved DT to VS. In addition, 17 PLs had higher YP than IR64 and the remaining 31 PLs had a similar YP IR64 under irrigated conditions. Detailed characterization of the PLs revealed three possible mechanisms that functioned together to contribute to their improved DT. The most important mechanism was dehydration avoidance (DA), characterized by significantly higher growth rate and biomass of all PLs than IR64 under stress and no reduction in biomass under control conditions. The second mechanism was efficient partitioning, characterized by improved harvest index in all PLs compared with IR64, resulting primarily from heavier grain weight and/or higher spikelet fertility under control conditions, which was the major constituent of the improved YP in the 17 best performing PLs. Drought escape (DE) by accelerated heading under drought was the third mechanism that contributed to DT of the PLs to RS. The considerable variation in the measured traits among the PLs with similar levels of DT and YP implies the complex genetic control of the mechanisms for DT/YP and offers opportunities to improve DT and YP further by fine-tuning of a small number of QTLs segregating among the PLs using MAS. Finally, our results indicate that selection for yield plus some secondary traits under appropriate type(s) of stress and nonstress conditions similar to the target environments are critically important for improving both DT and YP in rice.
Effect of genetic background on detection of quantitative trait locus (QTL) governing salinity to... more Effect of genetic background on detection of quantitative trait locus (QTL) governing salinity tolerance (ST) was studied using two sets of reciprocal introgression lines (ILs) derived from a cross between a moderately salinity tolerant japonica variety, Xiushui09 from China, and a drought tolerant but salinity susceptible indica breeding line, IR2061–520–6-9 from the Philippines. Salt toxicity symptoms (SST) on leaves, days to seedling survival (DSS), and sodium and potassium uptake by shoots were measured under salinity stress of 140 mmol/L of NaCl. A total of 47 QTLs, including 26 main-effect QTLs (M-QTLs) and 21 epistatic QTLs (E-QTLs), were identified from the two sets of reciprocal ILs. Among the 26 M-QTLs, only four (15.4%) were shared in the reciprocal backgrounds while no shared E-QTLs were detected, indicating that ST QTLs, especially E-QTLs, were very specific to the genetic background. Further, 78.6% of the M-QTLs for SST and DSS identified in the reciprocal ILs were als...
SummaryHybrid breeding is the main strategy for improving productivity in many crops, especially ... more SummaryHybrid breeding is the main strategy for improving productivity in many crops, especially in rice and maize. Genomic hybrid breeding is a technology that uses whole‐genome markers to predict future hybrids. Predicted superior hybrids are then field evaluated and released as new hybrid cultivars after their superior performances are confirmed. This will increase the opportunity of selecting true superior hybrids with minimum costs. Here, we used genomic best linear unbiased prediction to perform hybrid performance prediction using an existing rice population of 1495 hybrids. Replicated 10‐fold cross‐validations showed that the prediction abilities on ten agronomic traits ranged from 0.35 to 0.92. Using the 1495 rice hybrids as a training sample, we predicted six agronomic traits of 100 hybrids derived from half diallel crosses involving 21 parents that are different from the parents of the hybrids in the training sample. The prediction abilities were relatively high, varying f...
Hybrid rice varieties can outyield the best inbred varieties by 15 – 30% with appropriate managem... more Hybrid rice varieties can outyield the best inbred varieties by 15 – 30% with appropriate management. However, hybrid rice requires more inputs and management than inbred rice to realize a yield advantage in high-yielding environments. The development of stress-tolerant hybrid rice with lowered input requirements could increase hybrid rice yield relative to production costs. We used genomic prediction to evaluate the combining abilities of 564 stress-tolerant lines used to develop Green Super Rice with 13 male sterile lines of the International Rice Research Institute for yield-related traits. We also evaluated the performance of their F1 hybrids. We identified male sterile lines with good combining ability as well as F1 hybrids with potential further use in product development. For yield per plant, accuracies of genomic predictions of hybrid genetic values ranged from 0.490 to 0.822 in cross-validation if neither parent or up to both parents were included in the training set, and b...
The underground reserve (root) has been an uncharted research territory with its untapped genetic... more The underground reserve (root) has been an uncharted research territory with its untapped genetic variation yet to be exploited. Identifying ideal traits and breeding new rice varieties with efficient root system architecture (RSA) has great potential to increase resource-use efficiency and grain yield, especially under direct-seeded rice, by adapting to aerobic soil conditions. In this review, we tried to mine the available research information on the direct-seeded rice (DSR) root system to highlight the requirements of different root traits such as root architecture, length, number, density, thickness, diameter, and angle that play a pivotal role in determining the uptake of nutrients and moisture at different stages of plant growth. RSA also faces several stresses, due to excess or deficiency of moisture and nutrients, low or high temperature, or saline conditions. To counteract these hindrances, adaptation in response to stress becomes essential. Candidate genes such as early ro...
We studied variation in adaptive traits and genetic association to understand the low P responses... more We studied variation in adaptive traits and genetic association to understand the low P responses, including the symbiotic association of arbuscular mycorrhizal (AM) fungal colonization in Oryza species (O. sativa, O. nivara, and O. rufipogon). In the present experiment, we performed the phenotypic variability of the morphometric and geometric traits for P deficiency tolerance and conducted the association studies in GLM and MLM methods. A positive association between the geometric trait of the top-view area and root traits suggested the possibility of exploring a non-destructive approach in screening genotypes under low P. The AMOVA revealed a higher proportion of variation among the individuals as they belonged to different species of Oryza and the NM value was 2.0, indicating possible gene flow between populations. A sub-cluster with superior-performing accessions had a higher proportion of landraces (42.85%), and O. rufipogon (33.3%) was differentiated by four Pup1-specific mark...
Achieving sufficient food with finite resources and leaving lesser impact on the environment to f... more Achieving sufficient food with finite resources and leaving lesser impact on the environment to feed the growing global population has always been a great challenge. Here we review the concept and practices of Green Super Rice (GSR) that has led to a paradigm shift in the goals of crop genetic improvement and models of food production for promoting sustainable agriculture. The momentous achievements and global deliveries of GSR have been fueled through the integration of abundant genetic resources, functional gene discoveries, innovative breeding techniques with precise gene and whole-genome selections, and efficient agronomic management for promoting resource-saving and environment-friendly crop production systems. We also provide perspectives of new horizons in future genomic breeding technologies towards delivering green and nutritious crop varieties to further enhance the development of green agriculture to better nourish the world population.
Nitrogen is one of the most important nutrients for rice growth and development. Breeding of nitr... more Nitrogen is one of the most important nutrients for rice growth and development. Breeding of nitrogen deficiency tolerance (NDT) variety is considered to be the most economic measure to solve the constrain of low nitrogen stress on grain yield in rice. An interconnected breeding (IB) population of 497 lines developed using Huanghuazhan (HHZ) as the recurrent parent and eight elite lines as the donor parents were tested for five traits including grain yield, biomass, harvest index, thousand grain weight, and spikelet fertility under two nitrogen treatments in three growing seasons. Association analysis using 7,388 bins generated by sequencing identified a total of 14, 14, and 12 QTLs for the five traits under low nitrogen (LN), normal nitrogen (NN), and LN/NN conditions, respectively, across three seasons. Favorable alleles were dissected for the 40 QTLs at the 10 NDT regions, and OM1723 was considered as the most important parent with the highest frequency of favorable alleles contr...
Rice remains a major staple food source for the rapidly growing world population. However, regula... more Rice remains a major staple food source for the rapidly growing world population. However, regular occurrences of carcinogenic arsenic (As) minerals in waterlogged paddy topsoil pose a great threat to rice production and consumers across the globe. Although As contamination in rice has been well recognized over the past two decades, no suitable rice germplasm had been identified to exploit in adaptive breeding programs. Therefore, this current study identified suitable rice germplasm for As tolerance and exclusion based on a variety of traits and investigated the interlinkages of favorable traits during different growth stages. Fifty-three different genotypes were systematically evaluated for As tolerance and accumulation. A germination screening assay was carried out to identify the ability of individual germplasm to germinate under varying As stress. Seedling-stage screening was conducted in hydroponics under varying As stress to identify tolerant and excluder genotypes, and a fie...
BackgroundImproving performance of rice under drought stress has potential to significant impact ... more BackgroundImproving performance of rice under drought stress has potential to significant impact on rice productivity. Previously we reported that OsLG3 positively control rice grain length and yield.ResultsIn this study, we found that OsLG3 was more strongly expressed in upland rice compared to lowland rice under drought stress condition. Candidate gene association analysis showed that the natural variation in OsLG3 was associated with tolerance to water deficit stress in germinating rice seeds. Transgenic rice with enhanced OsLG3 expression exhibited improved tolerance to drought and that is most likely due to enhanced ROS scavenging efficiency. Phylogenetic analysis and pedigree records indicated that the tolerant allele of OsLG3 has potential to improve drought tolerance of japonica rice.ConclusionsCollectively, our work revealed that the natural variation of OsLG3 contributes to rice drought tolerance and the elite allele of OsLG3 is a promising genetic resource for the develop...
To develop superior rice varieties with improved yield in most rainfed areas of Asia/Africa, we s... more To develop superior rice varieties with improved yield in most rainfed areas of Asia/Africa, we started an introgression-breeding program for simultaneously improving yield and tolerances of multiple abiotic stresses. Using eight BC1 populations derived from a widely adaptable recipient and eight donors plus three rounds of phenotypic selection, we developed 496 introgression lines (ILs) with significantly higher yield under drought, salt and/or non-stress conditions in 5 years. Six new varieties were released in the Philippines and Pakistan and many more are being evaluated in multi-location yield trials for releasing in several countries. Marker-facilitated genetic characterization revealed three interesting aspects of the breeding procedure: (1) the donor introgression pattern in specific BC populations was characteristic; (2) introgression frequency in different genomic regions varied considerably, resulting primarily from strong selection for the target traits; and (3) signific...
Cold stress is one of the major abiotic stresses that impede rice production. A interconnected br... more Cold stress is one of the major abiotic stresses that impede rice production. A interconnected breeding (IB) population consisted of 497 advanced lines developed using HHZ as the recurrent parent and eight diverse elite indica lines as the donors were used to identify stably expressed QTLs for CT at the booting stage. A total of 41,754 high-quality SNPs were obtained through re-sequencing of the IB population. Phenotyping was conducted under field conditions in two years and three locations. Association analysis identified six QTLs for CT on the chromosomes 3, 4 and 12. QTL qCT-3-2 that showed stable CT across years and locations was fine-mapped to an approximately 192.9 kb region. Our results suggested that GWAS applied to an IB population allows better integration of gene discovery and breeding. QTLs can be mapped in high resolution and quickly utilized in breeding.
Interaction between rice (Oryza sativa L.) and its bacterial blight (BB) pathogen Xanthomonas ory... more Interaction between rice (Oryza sativa L.) and its bacterial blight (BB) pathogen Xanthomonas oryzae pv. oryzae (Xoo) is the model system between monocot plants and their bacterial pathogens. To understand the genome-wide interactions between Xoo and rice resulting from a broad-spectrum hypersensitive reaction (BSHR) mediated by a new rice resistance (R) gene, Xa39, comparative dynamic transcriptomic profiles in the incompatible and compatible interactions were investigated using three related rice lines and a highly virulent Xoo isolate, PXO99. Large numbers of rice and Xoo differentially expressed genes (DEGs) were identified in comparisons between the incompatible interactions and compatible ones, suggesting the gene network consisted of 27 genes in four groups of distinct functions involving leading to BSHR of rice in the sequential events from the avrXa39 ´ Xa39 interaction ® signal recognition and transduction ® protein modification ® programmed cell death. Correspondingly, several groups of Xoo genes expressed or upregulated specifically in the incompatible interaction were those for type III secretion system (T3SS) and type III secretion effectors (T3SEs). Combined evidence suggests LOC_Os11g37759, one of a two-member CC-NBS-LRR gene family on rice chromosome 11, as the most likely candidate for Xa39 in rice and two genes, XopN and XopX involved in T3SS of Xoo, as the most likely candidate genes for the corresponding avrXa39 in Xoo. Our transcriptome data and identified rice and Xoo DEGs provided a valuable source of information for future investigations on the Xoo-rice interactions. B acterial blight caused by Xanthomonas oryzae pv. oryzae (Xoo) is one of the most widely distributed rice diseases in tropical Asia and Africa, causing severe rice yield loses as most high-yielding rice cultivars are highly susceptible (NiÑo-Liu et al., 2006). Developing cultivars resistant to BB is the most economic and effective method to control the disease. Understanding of the molecular mechanisms underlying interactions between host and pathogen is required for successful knowledge-based rice improvement.
Background: Bacterial blight, caused by Xanthomonas oryzae pv. oryzae (Xoo), is a devastating ric... more Background: Bacterial blight, caused by Xanthomonas oryzae pv. oryzae (Xoo), is a devastating rice disease worldwide. Xa39 is a resistance (R) gene with a broad-spectrum hypersensitive response (BSHR) to Xoo. Nevertheless, the molecular mechanisms of resistance mediated by Xa39 remain unclear. In this study, the transcriptome profiling of a rice line carrying Xa39 and its parents at the early stage of Xoo infection were investigated. Results: A rice introgression line H471 carrying Xa39 exhibited a typical local hypersensitive response phenotype, accompanied by programmed cell death after inoculation with the Xoo Philippines' race 9b. Transcriptome profiling of H471 and its parents at 1 and 2 days post-inoculation was performed using RNA sequencing. In total, 306 differentially expressed genes (DEGs) were identified in H471 compared with its recurrent parent Huang-Hua-Zhan after inoculation with Xoo. Among them, 121 (39.5%) genes, with functional enrichments that were related to defense response, protein amino acid phosphorylation, and apoptosis, were found to be constitutively expressed. The other 185 (60.5%) genes, with GO terms that belonged to defense response, were significantly responsive to Xoo infection in H471. Ten up-regulated and 12 down-regulated genes encoding intracellular immune receptors were identified in H471 compared with Huang-Hua-Zhan. LOC_Os11g37759, which was located in the fine-mapping region harboring Xa39, is a Xa39 candidate gene. The putative BSHR-related co-regulatory networks were constructed using 33 DEGs from four functional groups, including gibberellic acid receptors and brassinosteroid regulators, which were differentially co-expressed with LOC_Os11g37759 in infected H471. Our results indicated that there might be cross-talk between the Xa39-mediated signal transduction cascades and the GA/BR signaling pathway, and that the defense mechanism was related to diverse kinases, transcription factors, post-translational regulation, and R genes. Conclusions: The present study provides the comprehensive transcriptome profile of a rice introgression line carrying Xa39 and its parents, and identifies a set of DEGs involved in BSHR mediated by Xa39. These data provide novel insights into the regulatory networks of plant disease resistance mediated by R genes, and the identified DEGs will serve as candidates for Xa39 cloning and for further understanding the molecular mechanism of BSHR.
ABSTRACT Four improved indica varieties adapted to high altitude regions namely K 39-2, lET 11988... more ABSTRACT Four improved indica varieties adapted to high altitude regions namely K 39-2, lET 11988, Himdhan and lET 11990 were treated with ethyl methane sulfonate, sodium azide and gamma rays with the objective of determining the most efficient mutagen and most responsive genotype for induction of mutations affecting predominantly reproductive parts and their functioning in rice. Frequency and spectrum of mutations affecting floral traits computed on Ml plant progeny and M2 population basis revealed the combination of the efficient sodium azide and the most responsive variety lET 11990 to be ideal for recovering highest number of selectively male sterile mutations. Among three kinds of male steriles viz., typical abortive, sporogenous and deformed male sex types recovered, critical study of abortive type helped identify several temperature sensitive genic male steriles, two of which are being improved for commercial exploitation in two-line hybrid breeding.
Proceedings of the National Academy of Sciences, 2006
Complete resistance (CR) and partial resistance (PR) of rice ( Oryza sativa L.) to its bacterial ... more Complete resistance (CR) and partial resistance (PR) of rice ( Oryza sativa L.) to its bacterial pathogen, Xanthomonas oryzae pv. oryzae ( Xoo ), was genetically dissected by using 2 mapping populations and 10 Xoo races. Two CR genes, 50 quantitative resistance loci, and 60 digenic interactions were identified, which showed various degrees of race specificity to the Xoo races. The complex epistasis between these loci led us to the discovery of complex genetic networks underlying the rice defensive system to Xoo . The networks consisted of two major components: one representing interactions between alleles at the R loci of rice and alleles at the corresponding avirulence loci of Xoo for CR and the other comprising interactions between quantitative resistance loci in rice and their corresponding aggressiveness loci in Xoo for PR. The race specificity of PR and its strong genetic overlap with CR indicate that PR is essentially “weaker” CR. The genetic networks discovered are expected t...
Great progress has been made in genetic dissection of quantitative trait variation during the pas... more Great progress has been made in genetic dissection of quantitative trait variation during the past two decades, but many studies still reveal only a small fraction of quantitative trait loci (QTLs), and epistasis remains elusive. We integrate contemporary knowledge of signal transduction pathways with principles of quantitative and population genetics to characterize genetic networks underlying complex traits, using a model founded upon one-way functional dependency of downstream genes on upstream regulators (the principle of hierarchy) and mutual functional dependency among related genes (functional genetic units, FGU). Both simulated and real data suggest that complementary epistasis contributes greatly to quantitative trait variation, and obscures the phenotypic effects of many 'downstream' loci in pathways. The mathematical relationships between the main effects and epistatic effects of genes acting at different levels of signaling pathways were established using the quantitative and population genetic parameters. Both loss of function and ''coadapted'' gene complexes formed by multiple alleles with differentiated functions (effects) are predicted to be frequent types of allelic diversity at loci that contribute to the genetic variation of complex traits in populations. Downstream FGUs appear to be more vulnerable to loss of function than their upstream regulators, but this vulnerability is apparently compensated by different FGUs of similar functions. Other predictions from the model may account for puzzling results regarding responses to selection, genotype by environment interaction, and the genetic basis of heterosis.
Three populations with a total of 125 BC2F3:4 introgression lines (ILs) selected for high yields ... more Three populations with a total of 125 BC2F3:4 introgression lines (ILs) selected for high yields from three BC2F2 populations were used for genetic dissection of rice yield and its related traits. The progeny testing in replicated phenotyping across two environments and genotyping with 140 polymorphic simple sequence repeat markers allowed the identification of 21 promising ILs that had significantly higher yields than the recurrent parent Shuhui527 (SH527). A total of 94 quantitative trait loci (QTL) were identified using the selective introgression method based on Chi-squared (χ2) and multi-locus probability tests and the RSTEP-LRT method based on stepwise regression. These QTL were mostly mapped to 12 clusters on seven rice chromosomes. Several important properties of the QTL affecting grain yield (GY) and its related traits were revealed. The first one was the presence of strong and frequent non-random associations between or among QTL that affect low-heritability traits (GY and spikelet number per panicle, SN) in the ILs with high trait values. Second, beneficial alleles at 88.9 % GY and 75 % SN QTL for increased productivity were from the donors, suggesting that direct phenotypic selection for high yield in our introgression breeding program was a powerful way to transfer beneficial alleles at many loci from the donors into SH527. Third, most QTL were in clusters with large effects on multiple traits, which should be the focal points in further investigations and marker-assisted selection in rice. The majority of the QTL identified were expressed only in one of the environments, suggesting that differential expression of QTL in different environments is the primary genetic basis of genotype × environment interaction. Finally, a large variation in both the direction and magnitude of QTL effects was detected for different donor alleles at seven QTL in the same genetic background and environments. This finding suggests the possible presence of functional diversity among the donor alleles at these loci. The promising ILs and QTL identified provide valuable materials and genetic information for further improving the yield potential of SH527, which is a backbone restorer of hybrid rice in China.
Drought is the most important factor limiting rice productivity in the rainfed areas of Asia. In ... more Drought is the most important factor limiting rice productivity in the rainfed areas of Asia. In this study, 48 pyramiding lines (PLs) and their recurrent parent, IR64, were evaluated over two years for their yield performances and related traits under severe drought stress at the reproductive stage (RS), the vegetative stage (VS) and irrigated control in order to understand the relationship between drought tolerance (DT) and yield potential (YP) in rice and their underlying mechanisms. When compared with IR64, all PLs had significantly improved DT to RS and 36 PLs also had significantly improved DT to VS. In addition, 17 PLs had higher YP than IR64 and the remaining 31 PLs had a similar YP IR64 under irrigated conditions. Detailed characterization of the PLs revealed three possible mechanisms that functioned together to contribute to their improved DT. The most important mechanism was dehydration avoidance (DA), characterized by significantly higher growth rate and biomass of all PLs than IR64 under stress and no reduction in biomass under control conditions. The second mechanism was efficient partitioning, characterized by improved harvest index in all PLs compared with IR64, resulting primarily from heavier grain weight and/or higher spikelet fertility under control conditions, which was the major constituent of the improved YP in the 17 best performing PLs. Drought escape (DE) by accelerated heading under drought was the third mechanism that contributed to DT of the PLs to RS. The considerable variation in the measured traits among the PLs with similar levels of DT and YP implies the complex genetic control of the mechanisms for DT/YP and offers opportunities to improve DT and YP further by fine-tuning of a small number of QTLs segregating among the PLs using MAS. Finally, our results indicate that selection for yield plus some secondary traits under appropriate type(s) of stress and nonstress conditions similar to the target environments are critically important for improving both DT and YP in rice.
Effect of genetic background on detection of quantitative trait locus (QTL) governing salinity to... more Effect of genetic background on detection of quantitative trait locus (QTL) governing salinity tolerance (ST) was studied using two sets of reciprocal introgression lines (ILs) derived from a cross between a moderately salinity tolerant japonica variety, Xiushui09 from China, and a drought tolerant but salinity susceptible indica breeding line, IR2061–520–6-9 from the Philippines. Salt toxicity symptoms (SST) on leaves, days to seedling survival (DSS), and sodium and potassium uptake by shoots were measured under salinity stress of 140 mmol/L of NaCl. A total of 47 QTLs, including 26 main-effect QTLs (M-QTLs) and 21 epistatic QTLs (E-QTLs), were identified from the two sets of reciprocal ILs. Among the 26 M-QTLs, only four (15.4%) were shared in the reciprocal backgrounds while no shared E-QTLs were detected, indicating that ST QTLs, especially E-QTLs, were very specific to the genetic background. Further, 78.6% of the M-QTLs for SST and DSS identified in the reciprocal ILs were als...
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Papers by Jauhar Ali