栽培稻与普通野生稻BC2F2群体产量相关性状的QTL分析

以广陆矮4号(Oryza sativa ssp. indica)为母本及轮回亲本,普通野生稻（Oryza rufipogon）为父本,分单株连续回交2次,构建BC2F2群体。首先用241对具有双亲多态性的SSR标记对BC2F1单株进行代换片段分析,在此基础上,根据BC2F1的表型选产量较优的单株自交获得BC2F2群体,用代换片段上具有双亲杂合型基因型的24对SSR标记进行QTL定位。在所选的BC2F1单株上,共检测到分布于7条染色体上的20个野生稻的代换片段,平均每条染色体上有2.86个;代换片段长度最小为0.55 cM,最大为33.00 cM,平均长度为12.36 cM,总覆盖长度为247.20 cM,覆盖率为16.21%。利用BC2F2群体对14个产量相关性状进行QTL定位,共检测到控制8个性状的20个QTL。对性状表型值起增效作用的有11个,占总检出数的55%。控制产量相关性状的QTL存在簇状分布现象,这与表型相关分析结果相符合。     

两个水稻抗褐飞虱隐性基因的遗传分析与初步定位

选用抗褐飞虱的海南普通野生稻渗入系WB01与感虫品种9311杂交,构建F2群体。采用141个具有多态性的SSR标记对303个F2:3株系进行分析,并应用MapMaker/EXP3.0和Windows QTL Cartographer2.0对水稻抗褐飞虱的数量性状基因座进行检测和遗传效应分析。共检测到2个抗性QTL,分别位于第4和第8染色体上,LOD值分别为2.92和3.15,贡献率分别为11.3%和14.9%。     

Yield Traits and Associated Marker Segregation in Elite Introgression Lines Derived from O. sativa × O. nivara

Introgression lines(ILs)derived from interspecific crosses are a source of new genetic variability.A total of 55 ILs derived from two crosses Swarna×O.nivara IRGC81848(population A)and Swarna×O.nivara IRGC81832(population B)were characterized for yield and yield-related traits/QTLs.Segregation of 103 simple sequence repeat(SSR)markers associated with yield-related QTLs was studied.Population A showed an average of 12.6%homozygous O.nivara alleles and population B showed 10.6%.Interestingly,three SSR markers,RM223,RM128 and RM517,showed conspicuous pattern of segregation.The distribution of parental alleles at three loci RM223,RM128 and RM517 linked to yield-related traits was unique.These markers flanked to several yield-related QTLs.RM223,flanking to qyld8.3,was heterozygous in almost all the 55 ILs except in IL10-3S and IL131S.RM128 on chromosome 1 and RM517 on chromosome 3 were mutually exclusive in 46 out of 55 ILs.These 46 ILs showed either of the marker allele RM128 or RM517 from O.nivara but not both.IL166S had both RM128and RM517 from O.nivara and the other ILs showed homozygous Swarna allele at RM517 except IL65S.Population structure assigned the 55 ILs to three sub-populations based on their genomic diversity.IL65S,IL166S,IL248S,IL7K and IL250K showed high yields in multi-location trials,and IL248S was released for cultivation as DRRDhan 40.     

Molecular Mapping of Sterility QTLs qSS-3, qSS-6a and qSS-7 as Single Mendelian Factors via NIL strategy

Hybrid sterility between Oryza glaberrima and O. sativa seriously hampers the introgression of favorable genes from each other. In order to further understand this issue, identification and isolation of hybrid sterility QTLs as single Mendelian factors are an effective strategy. A genetic map was constructed using a BC₁F₁ population derived from a cross between an O. sativa japonica cultivar and an O. glaberrima accession. Four main-effect QTLs for pollen sterility were detected in the BC₁F₁. Five BC₈F₁ advanced backcross populations were developed via successive backcrosses based on phenotype and molecular selections. The BC₈F₁ populations showed bimodal distribution for pollen fertility and could be classified into semi-sterile and fertile types, fitting single Mendilian factor inheritance ratios. Three QTLs detected in the BC₁F₁ corresponding to qSS-3, qSS-6a and qSS-7 were mapped on chromosomes 6, 3 and 7, respectively, as single Mendilian factors.     

小麦重要农艺性状QTL近等基因导入系的选育

为了给小麦重要农艺性状的QTL精细定位及克隆奠定基础,对14个从Am3/莱州953(轮回亲本)BC4F4代选出的性状表现与莱州953有明显差异的导入系的8个农艺性状进行了分析,结果表明,每个导入系有2～7个性状与莱州953差异显著,在每一个性状上都有对农艺性状具有正向效应的位点.利用143对在亲本之间具有多态性的SSR标记对导入系含有的供体片段进行了检测,其中54对引物在14个导入系中检测到了供体片段,每一个导入系中检测到3～15个纯合供体片段及0～4个杂合片段,占受体基因组的1.7%～14.2%,平均为7.48%.利用其中10个导入系与轮回亲本莱州953杂交的F2群体进行了单片段代换系的选择,从10个导入系的F2中检测到40个供体片段,并从F2群体中选出了22个单片段代换系.这些导入系及其单片段代换系可用于有益的QTL的发掘、QTL精细定位与作图等研究.     

Seed Length Controlled by Same Locus in Four Different AA GenomeSpecies of Genus Oryza简

To broaden the genetic basis and overcome the yield plateau in Asian cultivated rice, the exploitation and utilization of favorable alleles from rice species with the AA genome has become important and urgent in modern breeding programs. Four different interspecific populations were used to detect quantitative trait locus (QTL) for seed length, including a BC₄F₂ population derived from Oryza glumaepatula crossed with Dianjingyou 1 (a japonica cultivar), a BC₄F₂ population derived from O. nivara crossed with Dianjingyou 1, a BC₇F₁ population derived from a cross between O. longistaminata and RD23 (an indica cultivar), and a BC₈F₁ population derived from a cross between O. glaberrima and Dianjingyou 1. The QTLs for seed length in four different populations were termed as SL-3a, SL-3b, SL-3c and SL-3d, respectively. They had good collinearity and accounted for 49% to 60% of the phenotypic variations. Sequencing data indicated that four QTLs were different alleles of GS3 which were responsible for the seed length variation between O. sativa and its four AA genome relatives. These results will be valuable for confirming the evolution of GS3 and also be helpful for rice breeding.     

Development of elite breeding lines conferring Bph18 gene-derived resistance to brown planthopper (BPH) by marker-assisted selection and genome-wide background analysis in japonica rice (Oryza sativa L.)

Brown planthopper (BPH) is a serious threat to rice production. In this study, we have used the novel resistance gene Bph18 derived from Oryza australiensis and incorporated it into an elite japonica cultivar, Junambyeo, which is highly susceptible to BPH. The Bph18 gene was introduced by marker-assisted backcross (MAB) breeding into Junambyeo. The backcrossed progenies were evaluated for desirable agronomic and grain quality traits and the selection of improved breeding lines while simultaneously evaluating BPH resistance by bioassays in the greenhouse and foreground selection. Of the 26 advanced backcross breeding lines (ABL), four lines showed agronomic traits similar to those of the recurrent parent, with strong resistance to BPH. Molecular genotyping of the four ABL revealed the conversion of genotypes closely resembling the genotype of Junambyeo. The percentage of donor chromosome segments in ABL decreased from 12.3% in the BC₂ to 9.4%, 8.4% and 5.3% in BC₃, BC₄ and BC₅ generations, respectively. ABL retained small sizes of the donor chromosome segments on chromosomes 1, 2, 10, 11 and 12 but the genomes of ABL2, ABL3 and ABL4 were homosequential to the recurrent parent on chromosomes 3, 4, 5, 6, 7 and 9 without donor chromosome segment introgression. The ABL1 and ABL2 retained only some small segments of the donor genome on chromosomes 9 and 8, respectively. Fine structure analysis of the Bph18 flanking region between RM511 and RM1584 markers on chromosome 12 showed a progressive elimination of donor-derived chromosome segments from BC₂ to BC₅ generations. The percentage of O. australiensis derived chromosome segment substitution in the recurrent parent background decreased from 28% of the donor parent to 6.7%, 3.9%, 3.4% and 3.4% in BC₂, BC₃, BC₄ and BC₅ generations, respectively. However, it was revealed that the O. australiensis-derived chromosome segment (1320 kb) in ABL containing the Bph18 gene was consistently maintained irrespective of advances in backcross generations. BPH resistant elite breeding lines with agronomic and grain quality traits similar to those of the recurrent parent were successfully developed by foreground and background analysis in japonica background without linkage drag.     

Chromosome Segment Substitution Lines: A Powerful Tool for the Introgression of Valuable Genes from Oryza Wild Species into Cultivated Rice (O. sativa)

Wild species of rice (genus Oryza) contain many useful genes but a vast majority of these genes remain untapped to date because it is often difficult to transfer these genes into cultivated rice (Oryza sativa L.). Chromosome segment substitution lines (CSSLs) and backcross inbred lines (BILs) are powerful tools for identifying these naturally occurring, favorable alleles in unadapted germplasm. In this paper, we present an overview of the research involving CSSLs and BILs in the introgression of quantitative trait loci (QTLs) associated with the improved performance of rice including resistance to various biotic and abiotic stresses, and even high yield from wild relatives of rice and other unadapted germplasm into the genetic background of adapted rice cultivars. The CSSLs can be used to dissect quantitative traits into the component genetic factors and evaluate gene action as single factors (monogenic loci). CSSLs have the potential to uncover new alleles from the unadapted, non-productive wild rice accessions, develop genome-wide genetic stocks, and clone genes identified in QTL studies for functional genomics research. Recent development of high-density single-nucleotide polymorphism (SNP) arrays in rice and availability of custom-designed medium- and low-density SNP arrays will enhance the CSSL development process with smaller marker-defined segment introgressions from unadapted germplasm.     

Fine Mapping of QTLs for Stigma Exsertion Rate from Oryza glaberrima by Chromosome Segment Substitution

Stigma exsertion is an important trait for outcrossing ability in rice. Stigma exsertion rate (SER) of male sterile lines (MSLs) is a key factor affecting F₁-seed production in hybrid rice. In this study, seven QTLs for SER were detected on five chromosomes using a set of single-segment substitution lines (SSSLs) derived from O. glaberrima. Three of the QTLs were mapped in the estimated intervals of 92.5–333.0 kb. qSER-5 was located in a substitution segment of 92.5 kb. qSER-1b and qSER-8b were respectively limited to 333.0 kb and 107.5 kb by secondary substitution mapping. qSER-1b and qSER-3 had bigger additive effects of 11.5% and 11.9%, respectively, while the other five QTLs had smaller additive effects from 5.7% to 8.6%. Open reading frames were identified in the regions of qSER-5 and qSER-8b in O. sativa and O. glaberrima genomes. Fine mapping of the QTLs laid a foundation for the cloning of genes, and QTLs for SER will be used to develop MSLs with strong ability of outcrossing.     

Understanding Brown Planthopper Resistance in Rice: Genetics,Biochemical and Molecular Breeding Approaches

Brown planthopper (BPH, Nilaparvata lugens Stål) is the most devastating pest of rice in Asia and causes significant yield loss annually. Around 37 BPH resistance genes have been identified so far in indica, African rice varieties along with wild germplasms such as Oryza officinalis, O. minuta, O. nivara, O. punctata, O. rufipogon and O. latifolia. Genes/QTLs involved in BPH resistance, including Bph1, bph2/BPH26, Bph3, Bph6, bph7, BPH9, Bph12, Bph14, Bph15, Bph17, BPH18, bph19, Bph20, Bph21(t), Bph27, Bph27(t), Bph28(t), BPH29, QBph3, QBph4, QBph4.2, Bph30, Bph32, Bph33, Bph35 and Bph36, have been fine-mapped by different researchers across the globe. The majority of genes/QTLs are located on rice chromosomes 1, 3, 4, 6, 11 and 12. Rice plants respond to BPH attack by releasing various endogenous metabolites like proteinase inhibitors, callose, secondary metabolites (terpenes, alkaloids, flavonoid, etc.) and volatile compounds. Besides that, hormonal signal pathways mediating (antagonistic/synergistic) resistance responses in rice have been well studied. Marker-assisted breeding and genome editing techniques can also be adopted for improving resistance to novel BPH biotypes.     

应用DNA标记分析稻飞虱的抗性基因

简要地回顾了水稻抗飞虱的遗传位点定位和作图的新进展.来自具有不同基因组的野生稻渗入系的4个抗褐飞虱基因Bph 1、 bph 2、 bph 4和Bph 9,以及4个暂定名抗褐飞虱基因Bph 10(t)、bph 11(t)、bph 12(t)和Bph 13(t),目前已被定位于水稻12条染色体中的5条.其中,Bph 1、 bph 2、 Bph 9和Bph 10(t)在水稻第12染色体的长臂上形成1个连锁区段,位于bph 2位点附近约25 cM.检测出几个对田间抗性和杀卵作用有影响的QTL.抗白背飞虱基因Wbph 1、 Wbph 2和Wbph 6(t)已经或暂时定位了.粳稻中对白背飞虱具有杀卵抗性的QTL已进行了详细的分析,在第6染色体的短臂上检测到有效的QTL,在同一位点鉴定出1个显性的杀卵基因Ovc.在杀卵基因Ovc存在时,第1染色体上的1个QTL和第5染色体上的2个QTL增加白背飞虱的卵死亡率.用DNA标记进行QTL作图可以加深人们对作物抗虫性中复杂的生理和遗传机理的理解.标记辅助选择可以加速培育具多基因抗虫性的作物,还可以将野生种中的有利抗虫特性转入改良品种中,增加作物抗虫性的持久性和遗传多样性.     

Analysis of QTLs for seed low temperature germinability and anoxia germinability in rice (Oryza sativa L.)

Direct seeding instead of transplanting for rice (Oryza sativa L.) has increasingly been used in northern and eastern China because of labor and cost saving. However, poor germinability is still one of the major problems faced in the adoption of direct seeding under low temperature (low temperature germinability: LTG) and anoxia (anoxia germinability: AG) condition. To gain an understanding of the genetic control of seed germinability under these unfavourable conditions, two rice lines, USSR5 (japonica type) and N22 (indica type) and F₂ individuals derived from the cross USSR5 × N22 were tested for LTG and AG. USSR5 and N22 differ significantly for both LTG and AG. The LTG of the F₂ individuals ranged from 0 to 100% after a 10 days incubation. AG ranged from 0.0 to 4.0 cm shoot length. Based on segregation in the F₂ population, a linkage map was constructed using 121 SSR markers. The map covered 1821.5 cM, with a mean inter-marker distance of 16.7 cM. Eleven putative QTLs for LTG were detected, one on each of chromosomes 3–5, 7, 9–11, and four on chromosome 5. The USSR5 alleles in all these QTLs acted to increase LTG. Two QTLs for AG were located on chromosomes 5 and 11, respectively, at both of which the USSR5 alleles acted to increase AG. We propose that USSR5 could make a major contribution to improving LTG and AG in rice breeding programs.     

栽培稻与药用野生稻杂种后代的形态学和细胞遗传学研究

 选用两个抗褐飞虱的药用野生稻编号材料与两个感虫栽培品种杂交，通过胚培养获得了F₁、BC₁F₁和F₂植株。F₁杂种表现出野生亲本紫色柱头、长芒及抗褐飞虱等特性，其花粉母细胞中期I染色体多为单价体(仅0.77个二价体)，甚至不能形成花药，因而表现雄性不育，仅获得一株F₂，染色体数目为2n=46。用栽培稻回交获得BC₁F₁植株为异源三倍体，中期I染色体构型为12.25Ⅰ+11.85Ⅱ+0.01Ⅲ。     

高油玉米突变体子粒油分QTL定位

以EMS诱变获得的高油玉米突变体ce03005为材料,对子粒油分含量进行遗传分析,并以B73×ce03005的BC群体构建连锁图谱,考察玉米167个BC1S1家系的油分含量的变化。结果表明,利用101个SSR标记,所构建遗传图谱总长度为1 611.7 c M,标记间平均距离为15.9 c M。用复合区间作图法进行QTL分析,共检测到5个控制子粒油分含量的主效QTL和6个微效QTL,分别位于第1、6、7染色体上;所有QTL位点的含油量正向贡献均来自高油突变体;单个油分QTL的贡献率变幅为4.58%~18.62%。     

Localization,validation and characterization of plant-type QTLs on chromosomes 4 and 6 in rice (Oryza sativa L.)

Improvement of rice (Oryza sativa L.) plant type is a major breeding objective. This study aimed to precisely localize and characterize key genomic regions for plant type using near-isogenic individuals. Selfing of partially heterozygous F₅ recombinant inbred (RI) individuals [parental varieties Milyang 23 (M23) and Akihikari (AK)] developed heterogeneous inbred families (HIFs) composed of 108 and 93 F₇ progenies, which segregated at molecular marker loci on the long arms of chromosomes 4 and 6, respectively. The progeny lines (F₈) were evaluated for traits composing plant type in Los Baños, Philippines, to locate quantitative trait loci (QTLs) using interval mapping and to evaluate the effects of the QTL region by phenotypic comparison between the genotypes. QTLs for the traits were detected in 17 cM across XNpb12 on chromosome 6. The M23 homozygote for the QTL region resulted in a >7% increase and decrease in plant length and tiller number at heading, respectively, relative to the AK homozygote. Consequently, culm length (CL) and traits determining flag-leaf and panicle sizes increased 5–56% by the M23 homozygote, together with a 15% decrease in panicle number. For a QTL region detected in 6 cM across XNpb235 on chromosome 4, the AK homozygote had similar effects on these traits, except CL. The directions and magnitudes of their effects agreed with those previously estimated in the RI line population, thus increasing confidence in primary QTL analyses for plant type. Analyzing the HIFs validated and characterized the two QTL regions greatly involved in determining varietal plant type from an early growth stage to maturity, providing information useful for empirical and marker-assisted breeding towards rice improvement.     

Identification of QTLs for Improvement of Plant Type in Rice (Oryza sativa L.) Using Koshihikari / Kasalath Chromosome Segment Substitution Lines and Backcross Progeny F2 Population

Abstract

Thirty-nine chromosome segment substitution lines (CSSLs) population derived from a Koshihikari / Kasalath cross was used for quantitative trait locus (QTL) analysis of plant type in rice (Oryza sativa L.). Putative rough QTLs (26.2～60.3cM of Kasalath chromosomal segments) for culm length, plant height, panicle number, chlorophyll content of flag leaf blade at heading and specific leaf weight, were mapped on the several chromosomal segments based on the comparison of CSSLs with Koshihikari in the field experiment for 3 years. In order to verify and narrow QTLs detected in CSSLs, we conducted QTL analyses using F₂ populations derived from a cross between Koshihikari and target CSSL holding a putative rough QTL. The qPN-2, QTL for panicle number was mapped on chromosome 2. In traits of flag leaf, the qCHL-4-1 and qCHL-4-2 for chlorophyll content was mapped on chromosome 4, and the qSLW-7 for specific leaf weight on chromosome 7. All QTLs were detected in narrow marker intervals, compared with rough QTLs in CSSLs. The qPN-2, qCHL-4-1 and qCHL-4-2 had only additive effect. On the other hand, the qSLW-7 showed over-dominance. It could be emphasized that QTL analysis in the present study with the combination of CSSLs and backcross progeny F₂ population can not only verify the rough QTLs detected in CSSLs but also estimate allelic effects on the QTL.     

Genetic Analysis and Preliminary Mapping of Two Recessive Resistance Genes to Brown Planthopper,Nilaparvata lugens Stl in Rice

An F2 population derived from the cross of WB01, an introgression line resistant to brown planthopper (BPH) originated from Oryza rufipogon Griff. and a susceptible indica variety 9311, was developed for genetic analysis and gene mapping. The population with 303 F2:3 families was genotyped by 141 simple sequence repeat (SSR) markers and used for gene mapping. Two softwares, Mapmaker/Exp 3.0 and Windows QTL Cartographer V2.0 were applied to detect QTLs. Totally, two QTLs resistant to BPH, named temporarily a...     

Yield Traits and Associated Marker Segregation in Elite Introgression Lines Derived from O. sativa × O. nivara

Introgression lines (ILs) derived from interspecific crosses are a source of new genetic variability. A total of 55 ILs derived from two crosses Swarna × O. nivara IRGC81848 (population A) and Swarna × O. nivara IRGC81832 (population B) were characterized for yield and yield-related traits/QTLs. Segregation of 103 simple sequence repeat (SSR) markers associated with yield-related QTLs was studied. Population A showed an average of 12.6% homozygous O. nivara alleles and population B showed 10.6%. Interestingly, three SSR markers, RM223, RM128 and RM517, showed conspicuous pattern of segregation. The distribution of parental alleles at three loci RM223, RM128 and RM517 linked to yield-related traits was unique. These markers flanked to several yield-related QTLs. RM223, flanking to qyld8.3, was heterozygous in almost all the 55 ILs except in IL10-3S and IL131S. RM128 on chromosome 1 and RM517 on chromosome 3 were mutually exclusive in 46 out of 55 ILs. These 46 ILs showed either of the marker allele RM128 or RM517 from O. nivara but not both. IL166S had both RM128 and RM517 from O. nivara and the other ILs showed homozygous Swarna allele at RM517 except IL65S. Population structure assigned the 55 ILs to three sub-populations based on their genomic diversity. IL65S, IL166S, IL248S, IL7K and IL250K showed high yields in multi-location trials, and IL248S was released for cultivation as DRRDhan 40.     

Quantitative Trait Locus Mapping of High Photosynthetic Efficiency and Biomass in Oryza longistaminata

Photosynthetic efficiency, a key trait that determines yield potential in rice, is quantitatively regulated by multiple genes. Utilization of valuable genetic resources hidden in wild rice is an effective way to improve rice photosynthesis and yield potential. In this study, 152 backcross inbred lines derived from wild rice Oryza longistaminata were explored for QTL mapping of photosynthetic rate (Pn) and biomass (BM) in natural fields. Five novel QTLs for Pn and seven QTLs for BM or daily biomass (DBM) derived from O. longistaminata were identified. One of these QTLs, qPn8.1, could significantly improve Pn and was located in a 68-kb region containing only 11 candidate genes. Meanwhile, qBM1.1 and qDBM1.1 for BM and DBM on chromosome 1 were overlapped with qPn1.1 for Pn from 9311, and could affect both Pn and BM in natural fields. These QTLs identified in O. longistaminata may provide a novel alternative to explore new genes and resources for yield potentiality, highlighting the important role of wild rice in rice breeding programs.     

Mapping QTLs using a novel source of salinity tolerance from Hasawi and their interaction with environments in rice

Background

Salinity is one of the most severe and widespread abiotic stresses that affect rice production. The identification of major-effect quantitative trait loci (QTLs) for traits related to salinity tolerance and understanding of QTL × environment interactions (QEIs) can help in more precise and faster development of salinity-tolerant rice varieties through marker-assisted breeding. Recombinant inbred lines (RILs) derived from IR29/Hasawi (a novel source of salinity) were screened for salinity tolerance in the IRRI phytotron in the Philippines (E1) and in two other diverse environments in Senegal (E2) and Tanzania (E3). QTLs were mapped for traits related to salinity tolerance at the seedling stage.

Results

The RILs were genotyped using 194 polymorphic SNPs (single nucleotide polymorphisms). After removing segregation distortion markers (SDM), a total of 145 and 135 SNPs were used to construct a genetic linkage map with a length of 1655 and 1662 cM, with an average marker density of 11.4 cM in E1 and 12.3 cM in E2 and E3, respectively. A total of 34 QTLs were identified on 10 chromosomes for five traits using ICIM-ADD and segregation distortion locus (SDL) mapping (IM-ADD) under salinity stress across environments. Eight major genomic regions on chromosome 1 between 170 and 175 cM (qSES1.3, qSES1.4, qSL1.2, qSL1.3, qRL1.1, qRL1.2, qFWsht1.2, qDWsht1.2), chromosome 4 at 32 cM (qSES4.1, qFWsht4.2, qDWsht4.2), chromosome 6 at 115 cM (qFWsht6.1, qDWsht6.1), chromosome 8 at 105 cM (qFWsht8.1, qDWsht8.1), and chromosome 12 at 78 cM (qFWsht12.1, qDWsht12.1) have co-localized QTLs for the multiple traits that might be governing seedling stage salinity tolerance through multiple traits in different phenotyping environments, thus suggesting these as hot spots for tolerance of salinity. Forty-nine and 30 significant pair-wise epistatic interactions were detected between QTL-linked and QTL-unlinked regions using single-environment and multi-environment analyses.

Conclusions

The identification of genomic regions for salinity tolerance in the RILs showed that Hasawi possesses alleles that are novel for salinity tolerance. The common regions for the multiple QTLs across environments as co-localized regions on chromosomes 1, 4, 6, 8, and 12 could be due to linkage or pleiotropic effect, which might be helpful for multiple QTL introgression for marker-assisted breeding programs to improve the salinity tolerance of adaptive and popular but otherwise salinity-sensitive rice varieties.