Dissection of component QTL expression in yield formation in rice

Yield and its components were investigated by using a population of 241 recombinant inbred lines (F₉ RILs) derived from an elite hybrid rice cross of ‘Zhenshan 97’בMinghui 63′. Quantitative trait loci (QTLs) for causal analysing of yield traits were detected at different yield component (YC) influences by conditional and unconditional QTL mapping methods. The number of QTLs significantly affecting yield was different at component‐special influence. Some QTLs controlling yield identified in one component influence were undetectable at the others. More QTLs for yield could be detected at different YC influences. It is possible to reveal that causal gene expression for yield could be different at different YC influences. Mapping QTLs for component effects of yield could help us in understanding the nature of cause‐effect traits for the formation of grain yield.     

Detection of quantitative trait locus for leaffolder (Cnaphalocrocis medinalis (Guenée)) resistance in rice on linkage group 1 based on damage score and flag leaf width

Rice leaffolder (RLF) (Cnaphalocrocis medinalis (Guenée) is a destructive and widespread insect pest throughout the rice growing regions in Asia. The genetics of resistance to RLF in rice is very complex and not thoroughly explored. The present study was conducted to detect the quantitative trait loci (QTL) associated with RLF resistance involving 176 recombinant inbred lines (RILs) of F₈ generation derived from a cross between IR36, a leaffolder susceptible variety and TNAULFR831311, a moderately resistant indica rice culture. Simple sequence repeat (SSR) markers were used to construct specific linkage groups of rice. All the RILs were screened to assess their level of resistance to RLF by measuring the leaf area damaged. Besides this, the length and width of the flag leaf of each RIL were measured since these two parameters were considered as correlated traits to the RLF resistance in rice. All the above parameters observed across the RILs showed quantitative variation. Correlation analysis revealed that damage score based on greenhouse screening was positively correlated with length and width of the flag leaf. Out of 364 SSR markers analysed, 90 were polymorphic between the parents. Multi-point analysis carried out on segregating 69 SSR marker loci linkage group wise resulted in construction of linkage map with eleven groups of 42 SSR markers. Through single marker analysis, 19 SSR markers were found to have putative association with the three phenotypic traits studied. Of these markers, RM472 was identified as a locus having major effect on RLF resistance trait based on length of the flag leaf. Interval mapping detected two QTLs on linkage group 1. Among these QTLs, the QTL flanked by RM576–RM3412 were found to be associated with width of the flag leaf and RLF resistance. The putative SSR markers associated with leaffolder resistance identified in the present study may be one of the loci contributing resistance to RLF in rice.     

水稻第1染色体长臂上微效千粒重QTL qTGW1.2的验证与分解

本文报道了水稻第1染色体长臂上微效千粒重QTL qTGW1.2的验证和分解。针对前期qTGW1.2定位结果, 应用SSR标记检测, 从籼籼交组合珍汕97³/密阳46衍生的1个BC₂F_{7分离群体中, 筛选到杂合区间分别为RM11621-RM297和RM212-RM265的2个单株, 构建了两套BC2F8:9近等基因系, 将qTGW1.2进一步界定在RM212-RM265及其两侧交换区间的区域内。}在此基础上, 筛选出5个在目标区间内分离片段缩小且呈阶梯状排列的单株, 衍生了5套BC₂F₁₀分离群体, 应用Windows QTL Cartographer 2.5进行QTL分析。结果表明, 每套群体均检测到千粒重QTL, 加性效应为0.13~0.38 g, 来自密阳46的等位基因提高千粒重; 经比较各个群体的分离区间, 将qTGW1.2分解为互引连锁的2个QTL, 其中, qTGW1.2a位于RM11730和RM11762之间934 kb的区域内, 呈加性作用, qTGW1.2b位于RM11800和RM11885之间2.1 Mb的区域内, 呈正向超显性。     

Novel quantitative trait loci for yield and yield related traits identified in Basmati rice (Oryza sativa)

Increasing crop productivity is one of the prime goals of crop breeding research. Rice grain yield is a complex quantitative trait governed by polygenes. Although several QTLs governing grain yield traits have been reported and limited attempts have been made to map QTLs for grain yield parameters in Basmati rice. A population from the cross Sonasal and Pusa Basmati 1121 comprising 352 RILs was generated through the single seed descent method. A total of 12 QTLs governing yield and yield-related traits were mapped on six chromosomes, namely, 1, 2, 3, 7, 8 and 9, of which five QTLs were novel. We identified a novel and robust epistatic QTL (qPH1.1 and qPL1.1) governing plant height and panicle length, flanked by the markers RM5336-RM1 on chromosome 1. The gene encoding brassinosteroid insensitive 1-associated receptor kinase 1 precursor is the putative candidate gene underlying this epistatic QTL. Another novel QTL, qNT3.1, governing tiller number was bracketed to a region of .77 Mb between the markers RM15247 and RM15281 on chromosome 3. Of the 57 annotated gene models, Os03g0437600 encoding alpha/beta-fold hydrolase, a homologous to AtKai2 is a putative candidate gene underlying the novel QTL qNT3.1. The other QTLs such as qDFF1.1 governing days to 50% flowering co-localizes with the gene Ghd7, QTL for plant height qPH1.2 co-localizes with the gene sd1, the QTLs for panicle length co-localizes with FUWA and DEP2, the QTL for tiller number co-localizes with OsRLCK57 and QTLs for thousand-grain weight co-localize with the major gene GS3. The QTLs identified in the current study can be effectively used in marker-assisted selection for developing Basmati rice varieties with a higher yield.     

Quantitative trait loci and candidate genes for yield and related traits in Madhukar x Swarna RIL population of rice

Yield of popular rice varieties such as Swarna grown in rainfed lowlands and Madhukar grown in flood prone areas needs to be continuously improved. Recombinant inbred lines (RILs) were developed from the cross between two indica cultivars Madhukar and Swarna. QTLs were mapped using 110 markers in 168 RILs. In all, 26 QTLs were mapped for yield and five related traits on chromosomes 1, 2, 3, 6, 7, 8, 10, 11, and 12. QTL for plant height and days to flowering were co-located between RM23147 — RM337 on chromosome 8. RM251, RM314, and RM1135 were significantly associated with plant height and OsYSL17 was significantly linked with grain yield. Epistatic interaction was detected for plant height and number of tillers. Several candidate genes reported for yield and related traits underlie the QTL regions.     

Identification and validation of a yield-enhancing QTL cluster in rice (Oryza sativa L.)

Improvement of rice grain yield (YD) is an important goal in rice breeding. YD is determined by its related traits such as spikelet fertility (SF), 1,000-grain weight (TGW), and the number of spikelets per panicle (SPP). We previously mapped quantitative trait loci (QTLs) for SPP and TGW using the recombinant inbred lines (RILs) derived from the crosses between Minghui 63 and Teqing. In this study, four QTLs for SF and four QTLs for YD were detected in the RILs. Comparison of the locations of QTLs for these three yield-related traits identified one QTL cluster in the interval between RM3400 and RM3646 on chromosome 3. The QTL cluster contained three QTLs, SPP3a, SF3 and TGW3a, but no YD QTL was located there. To validate the QTL cluster, a BC₄F₂ population was obtained, in which SPP3a, SF3 and TGW3a were simultaneously mapped to the same region. SPP3a, SF3 and TGW3a explained 36.3, 29.5 and 59.0 % of phenotype variance with additive effect of 16.4 spikelets, 6 % SF and 1.8 g grain weight, respectively. In the BC₄F₂ population, though the region has opposite effects on TGW and SPP/SF, a YD QTL YD3 identified in this cluster region can increase 4.6 g grains per plant, which suggests this QTL cluster is a yield-enhancing QTL cluster and can be targeted to improve rice yield by marker aided selection.     

Fine mapping of a major quantitative trait locus, <Emphasis Type="Italic">qFLL6.2</Emphasis>, controlling flag leaf length and yield traits in rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

Fine mapping of a quantitative trait locus, qFLL6.2, controlling flag leaf length (FLL) and yield traits in rice was conducted using four sets of near isogenic lines (NILs) that were developed from a common residual heterozygote at F₇ generation of the indica rice cross Zhenshan 97/Milyang 46. Each of the NIL sets consisted of 40 lines that are S₁ progenies of ten maternal homozygotes, ten paternal homozygotes, and 20 heterozygotes differing in a portion of the 1.19-Mb interval RM3414–RM6917 on the short arm of rice chromosome 6. Analysis of phenotypic differences among the three genotypic groups in each NIL set delimited qFLL6.2 to a 62.1-kb region flanked by simple sequence repeat marker RM3414 and sequence-tagged site marker Si2944. This QTL explained 52.73% of the phenotypic variance, and the Zhenshan 97 allele increased FLL by 2.40 cm. Based on data collected from homozygous lines of three of the NIL sets, qFLL6.2 was shown to have major effects on all the three yield traits analyzed, including the number of spikelets per panicle, the number of filled grains per panicle, and grain weight per panicle. A comparison of the different groups revealed that the effect of qFLL6.2 was highly consistent across different genetic backgrounds and environments, providing a good candidate for map-based cloning and investigating the source–sink relationship in rice.     

利用重组自交系群体对水稻产量相关性状的QTL分析

利用水稻(Oryza sativa L．)珍汕97和SLG组合的重组自交系群体和177个微卫星标记,采用复合区间作图法对产量相关性状进行了数量性状基因(QTL)定位,共得到38个QTLs。单个QTL解释这些性状6．6％～32．9％的性状变异,各性状QTL的累积表型贡献率达21．3％～79．5％。大多数性状之间具有显著的表型相关性,相关性较高的性状之间常具有较多共同或紧密连锁的QTL。     

Comparison of QTLs mapped in RILs and their test-cross progenies of tropical maize for insect resistance and agronomic traits

Quantitative trait loci (QTL) affecting resistance to south-western corn borer Diatraea grandiosella (SWCB) and sugarcane borer Diatraea saccharalis (SCB) have been identified previously in F_2:3 lines and recombinant inbred lines (RILs) of tropical maize using restriction fragment length polymorphism (RFLP) analyses. Our objective was to determine whether QTLs identified in these generations are also expressed in test crosses (TC) of RILs. A population of 166 TC progenies was developed by crossing RILs from the cross CML131 (susceptible) × CML67 (resistant) with the unrelated, susceptible tester line CML216. Resistance to first-generation SWCB, measured as leaf-feeding damage (LFD) under artificial infestation, and other agronomic traits were evaluated in two environments for the TC progenies and three environments for 183 RILs. The correlation between line per se and TC performance was low for LFD and intermediate for most agronomic traits. Estimates of the genotypic variance and heritabilities were smaller in the TC progenies than in the RILs for all traits. Quantitative trait loci were identified using an RFLP linkage map with 136 loci. For LFD, four QTLs were detected in the TC progenies, of which two were in common with nine QTLs previously mapped in the RILs. Few QTLs for agronomic traits were common to the two types of progeny, because of the low consistency of QTL positions for all traits in RIL and TC progenies, the use of TC progenies should be considered in QTL mapping studies as the first step for marker-assisted selection in hybrid breeding.     

Comparison of quantitative trait loci for 1,000-grain weight and spikelets per panicle across three connected rice populations

The ability to detect quantitative trait loci (QTLs) in a bi-allelic population is often limited. The power of QTL detection and identification of the most beneficial allele at each QTL could be greatly improved by comparing QTLs among different populations derived from connecting multi-parents. In this study, three sets of connected recombinant inbred lines (RILs) derived from the crosses between Zhenshan 97 and Minghui 63 (PZM), Zhenshan 97 and Teqing (PZT), and Minghui 63 and Teqing (PMT), respectively, were used. QTL analyses for the number of spikelets per panicle (SPP) and 1,000-grain weight (TGW) were performed in PZT, and five SPP QTLs on chromosomes 1, 6, and 7 and two TGW QTLs on chromosome 1 were detected. QTL for SPP was also identified in PMT, and six QTLs were detected on chromosomes 1, 2, 3, 6, and 7 in this population. In an earlier study, we identified five SPP QTLs and four TGW QTLs in PMT and nine TGW QTLs in PZM. Comparison of the QTL mapping results of these two studies showed that one QTL was common to the three populations, 11 QTLs were detected in two populations, and six QTLs were found in only one population. Comparison of genetic effect and the action direction of the QTLs detected in the three populations showed that additive effects of QTLs estimated in different populations were also expressed additively among three parental alleles. Additive effects of SPP7a estimated in three near-isogenic line F₂ populations supported this finding. Based on these results, we suggest that pyramiding the most beneficial alleles among the three parents could efficiently improve rice yield.     

水稻剑叶角度与主穗产量的遗传剖析

理想水稻株型的选育与高产育种密切相关,而剑叶角度则是构成水稻理想株型的重要指标之一,同时也是影响水稻产量的重要因素。合理开发利用水稻中控制剑叶角度及产量相关的数量性状基因座位（QTL）,并结合分子育种技术,可更好地为高产制繁种目标服务。通过应用由244个株系组成的珍汕97B/密阳46重组自交系（RIL）群体,构建含256个分子标记的连锁图谱,采用QTL区间作图法对剑叶角度及主穗产量等5个性状进行定位分析,共检测到17个QTL,分布于染色体1、2、3、5、6、9、10、11。这些QTL对相应性状的贡献率介于3.46%~25.64%之间。在第1染色体上检测到控制5个性状的QTL,其中控制剑叶角度的两个QTL;在第2、3、9、10、11染色体上分别检测到各一个QTL;第5染色体上检测到控制剑叶、每穗总粒数和每穗实粒数的3个QTL;1个每穗实粒数和2个每穗实粒重的QTL分布于第6染色体上。多个区间表现出对两个性状的显著作用,其中第1染色体2个,第6染色体1个。相关性分析表明,较小的剑叶角度可通过提高结实率进而显著增加产量。     

Genetic analysis and map-based delimitation of a major locus qSS3 for seed size in soybean

Seed size is a major determinant of grain yield in soybean, however their genetic basis remains largely unknown. In order to delimit map-based position of a major locus qSS3 , we evaluated three mapping populations, including RILs, NILs and a sub-F₂ in three environments for six seed size-related traits. For these traits, the kurtosis and skewness ranged between 0.0 and 1.16, while h²_b ranged from 0.75 to 0.96, indicating that this RIL population is suitable for QTL analysis. QTL analysis identified 12 loci which consist of 30 significant QTLs with PVE% and LOD values of 6.6%–26.2% and 2.50–5.61, respectively. Among them, qSS3 was a major and stable locus explaining 7.3%–26.2% of the variation in 5 of the 6 traits, with the respective LOD values falling in the range of 2.72–5.61. Additionally, qSS3 effects were confirmed in NILs and delimited to an interval of ~1,126 kb containing 123 annotated genes. Overall, this study may assist efforts aiming to improve soybean seed traits by identifying valuable genetic resources which can be used in future MAS breeding programmes.     

QTL analysis of cooked rice grain elongation, volume expansion, and water absorption using a recombinant inbred population

The traits of elongation, volume expansion, and water absorption are very important in determining the quality of cooked rice grains. In this study, quantitative trait loci (QTL) analysis of these traits was performed using a recombinant inbred population derived from a cross between two indica cultivars, ‘Zhenshan 97’ and ‘Minghui 63 ,’ which are the parents of the most widely grown hybrid rice in China. Using a linkage map based on 221 molecular marker loci covering a total of 1796 cM, a total of 33 QTLs were identified for the nine traits tested. QTLs were detected on chromosomes 1– 3 , 5– 9 , and 11 , respectively. The QTLs identified included three for cooked rice grain length elongation (chromosomes 2 , 6 , and 11), six for width expansion (chromosomes 1‐ 3 , 6 , 9 , and 11) and two for water absorption (chromosomes 2 and 6). Interestingly, a single QTL located near the wx gene on chromosome 6 seemed to influence all the traits tested for the cooked rice quality.     

QTLs identification of crude fat content in brown rice and its genetic basis analysis using DH and two backcross populations

Fat content is a concern for the enhancement of rice for eating, cooking, and storage qualities. To clarify its genetic mechanism, a double haploid (DH) population derived from anther hybrid F₁ of Zhenshan 97B (indica) and Wuyujing 2 (japonica) and two backcross F₁ (BCF₁) populations, which came from the DH lines backcrossing to two parents, were used to scan quantitative trait loci (QTLs) and dissect gene effects for the crude fat content (CFC) in brown rice. Fourteen QTLs were resolved, distributing on chromosomes 1, 3, and 5–9. Three loci were detected repeatedly in two populations, DH or BCF₁. Among these loci, a major QTL, qCFC5, flanking markers RM87 and RM334, was located on chromosome 5, which was detected simultaneously among three populations. The main QTLs had a major role in controlling CFC in brown rice and were modified by several mini-effect QTLs and epistatic affection. Wenjun Liu and Jing Zeng are contributed equally to this paper.     

Identification and application of major quantitative trait loci for panicle length in rice (Oryza sativa) through single‐segment substitution lines

Panicle length (PL), an important yield‐related trait, strongly affects yield components, such as grain number, grain density and rice quality. More than 200 panicle length quantitative trait loci (PL QTLs) are identified, but only a small number are applied in rice breeding. In this study, we performed QTL analysis for PL using 42 single‐segment substitution lines (SSSLs) derived from nine donors in the genetic background of HJX74. Fourteen QTLs and five heterosis QTLs (HQTLs) for PL were recognised. Three QTLs and four HQTLs acted positively, and the other eleven QTLs and one HQTL acted negatively. By scanning the single heterozygous background region of the F₂ population with large‐genetic‐effect SSSLs, we mapped PL loci qPL6‐2 and qPL7‐1 to different locations on chromosomes 6 and 7, respectively, in three consecutive years of independent trials. The genetic effects of these QTLs were further assessed. qPL6‐2 demonstrated the most positive additive effect (QTL), whereas qPL7‐1 achieved the most positive dominant effect (HQTL) for PL. These results indicated that the pyramiding of PL QTLs might increase grain yield and facilitate the application of the beneficial allele in hybrid rice breeding.     

稻米淀粉黏滞性QTL定位及其G×E互作分析

用珍汕97B/密阳46构建RIL群体及其遗传图谱,经海南和杭州两地遗传试验,以精米粉RVA谱5个参数特征值PKV、HPV、CPV、BDV和SBV作为研究稻米淀粉黏滞性的指标,运用检测QTL主效应、上位性效应和G×E互作效应的遗传分析方法,进行QTL联合分析。结果表明, (1)在检测到涉及5个性状的9个主效应QTL中,除PKV位于第5染色体qPKV5外,其余8个QTL均位于第6染色体上;(2)5个性状均检测到位于第6染色体RM197-RZ516区间的主效应QTL,很可能它们为同一基因,该基因还与Wx基因处于相同区域;(3)检测到与PKV、HPV、CPV、BDV等4个性状有关的QTL主效基因均表现有G×E互作,且方向一致,在海南试验中有增效作用;(4)还检测到涉及5个性状的10对上位性互作效应,但均未发现有显著的上位性×环境互作效应。     

应用重组自交系群体检测控制水稻糙米粗蛋白和粗脂肪含量的QTL

用协青早B/密阳46重组自交系群体及其分子连锁图谱,及Windows QTL Cartographer 2.0的复合区间作图法和多区间作图法,对水稻糙米蛋白质含量和粗脂肪含量进行QTL分析。检测到控制蛋白质含量的QTLs 5个（qPc-3、qPc-4、qPc-5、qPc-6、qPc-10）,单个QTL对群体表型变异的贡献率     

Genetic studies of seed longevity in hexaploid wheat using segregation and association mapping approaches

Xieyou9308 is the first commercial super hybrid rice released in 1996 in China. To clarify its genetic mechanism underlying high yield potential, a recombinant inbred line (RIL) population derived from the cross between the maintainer line XieqingzaoB (XQZB) and the restorer line Zhonghui9308 (ZH9308) and two derived backcross F₁ (BCF₁) populations were developed for the identification of quantitative trait loci (QTLs) related to ten important agronomic traits (tiller number (TN), heading date (HD), and grain yield per plant (GYPP), etc.). The BCF₁ performance was closely correlated with the performance of their parental RILs according to both the analysis of broad-sense heritability (h _B ²) and phenotypic correlation coefficient (PCC) in the two BCF₁ populations, but not proved by QTL analysis. A total of 21 additive-effect main QTLs (M-QTLs), 22 dominant-effect M-QTLs, and 19 dominant-effect M-QTLs were detected with the WinQTLCart 2.50 software for the ten traits in the RIL and two BCF₁ populations, respectively. Of theses, three QTLs (qHD7a, qPPP3a, and qPL10) of 21 were detected repeatedly in the RIL and one BCF₁ populations, ten QTLs underlying four traits were only detected repeatedly in two BCF₁ populations, and nine QTLs controlling more than two traits were detected repeatedly, the additive-effect QTLs and dominant-effect QTLs play an important role in the performance of agronomic traits and no epistatic QTL of additive by additive effect and dominant by dominant-effect was detected for all traits in three populations. This research is valuable for M-QTL related to important agronomic trait in future fine mapping and positional cloning.     

Identification and validation of a major QTL for salt tolerance in soybean

To identify quantitative trait loci (QTLs) conditioning salt tolerance in soybean (Glycine max (L.) Merr.), two recombinant inbred line (RIL) populations derived from crosses of FT-Abyara × C01 and Jin dou No. 6 × 0197 were used in this study. The FT-Abyara × C01 population consisted of 96 F₇ RILs, and the Jin dou No. 6 × 0197 population included 81 F₆ RILs. The salt tolerant parents FT-Abyara and Jin dou No. 6 were originally from Brazil and China, respectively. The QTL analysis identified a major salt-tolerant QTL in molecular linkage group N, which accounted for 44.0 and 47.1% of the total variation for salt tolerance, in the two populations. In the FT-Abyara × C01 population, three RILs were found to be heterozygous around the detected QTL region. By selfing the three residual heterozygous lines, three sets of near isogenic lines (NILs) for salt tolerance were developed. An evaluation of salt tolerance of the NILs revealed that all the lines with FT-Abyara chromosome segment at the QTL region showed significantly higher salt tolerance than the lines without the FT-Abyara chromosome segment. Results of the NILs validated the salt tolerance QTL detected in the RIL populations.     

Impact of seasonal changes on spikelets per panicle,panicle length and plant height in rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

Temperature and photoperiod fluctuate rapidly in different seasons of the year, and analyzing their effects on rice yield components is crucial for adaptation of rice under various climatic conditions. To study the effects of seasonal changes on yield components, 168 recombinant inbred lines derived from a cross between two Oryza sativa L. indica varieties, Zhenshan 97 and Zhongzao 18 were grown for phenotype collection, in three different seasons, within a year. The results implied that temperatures across the three seasons played a crucial role in determining the trait effects. Spikelets per panicle (SPP), panicle length (PL) and plant height (PH) traits increased with high temperatures in middle season. Genetic analysis detected major quantitative trait loci (QTLs) qSPP10, qPL10 and qPH10 for SPP, PL and PH in the interval between markers RM1375 and RM3229 on chromosome 10, in all the three seasons. Two-way ANOVA showed that genotype by environment and QTL by environment interactions for these traits were highly significant (P < 0.0001). The region with a cluster of QTLs detected in all three seasons could be the preferred target to breeders in developing rice varieties that can be accustomed to different seasonal changes.