Identification of QTLs for Cooking and Eating Quality of Rice Grain

Rice is one of the major crops served as the staple food of more than 50% of the world’s population. Recently, more attention has been paid to rice quality, especially the cooking and eating quality than achieving the higher yield. The amylose content (AC), gelatinization temperature (GT) and gel consistency (GC) are the three major rice characteristics directly related to cooking and eating quality, responsible for the physical and chemical characteristics of the starch in the endosperm [1…     

水稻粒重及其相关性状的遗传解析

 应用292个 Lemont/ 特青F13重组自交系(RILs)和272个标记的遗传连锁图谱分析粒重及籽粒长、宽、厚、长/宽、体积和容重6个相关性状的遗传。所有性状在RILs中出现超亲分离。粒重与其他籽粒性状均呈显著正相关，而涉及籽粒品质的容重与粒宽、粒厚呈显著负相关。检测到影响粒重及其相关性状的主效QTL 48个和互作位点38对，这些QTL解释了各性状55%以上的表型变异。80%以上的主效QTL呈现一因多效或生理相关。QGl3和 QGl2等通过籽粒体积间接影响粒重， Lemont增效等位基因有利于籽粒品质的改良，而QGw5、QGt1和QGv7的特青增效等位基因增加粒重同时降低籽粒容重及品质。通过标记辅助选择进行不同QTL的重组，有望在增加粒重的同时改善籽粒品质。     

QTL mapping of grain quality traits in rice

Grain quality improvement is one of the most important goals in a rice breeding program. An indica variety with small grain size was crossed to a japonica variety with large grain size to construct a set of recombinant inbred lines (RILs) which was used to identify quantitative trait loci (QTLs) controlling eight grain quality traits. Based on a linkage map of 185 SSR markers, a total of 16 QTLs were mapped on six chromosomes. A pleiotropic main effect QTL (M-QTL) flanked by RM3204 and RM16 on chromosome 3 influences the grain length (GL), length width ratio (LWR) and head rice ratio (HRR), explaining the phenotypic variation of 46.0, 36.1 and 29.7%, respectively. A total of 18 epistatic QTLs were identified for all the traits except MRR, distributed on all the chromosomes except chromosome 10. Two M-QTLs for GL and one M-QTL for GW were involved in epistatic QTL. No significant interaction between M-QTL or epistatic QTL and environment was detected except AC having significant M-QTL by environment interaction with minor effect. GL and LWR have a significant negative relation with HRR which might make it difficult to develop long grain with higher HRR in the rice breeding practice.     

多环境下粳稻产量及其相关性状的条件和非条件QTL定位

 为了剖析粳稻产量及其相关性状的遗传基础,利用粳稻品种秀水79×Ｃ堡衍生的重组自交系群体,在3个环境下对全生育期、株高、单株穗数、每穗粒数、百粒重、籽粒产量和生物产量进行了非条件和条件QTL定位。共检测到43个主效QTL和29对上位性QTL。利用非条件QTL定位方法检测到37个主效QTL和26对上位性QTL。其中,籽粒产量定位到3个主效QTL qGY1.2、qGY7.1和qGY9,未检测到上位性QTL。利用条件QTL方法分别将全生育期、株高、穗数、每穗粒数、百粒重和生物产量各自调整到同一水平后,籽粒产量共检测到9个主效条件QTL和3对上位性QTL,其中3个主效QTL与非条件下定位到的相同。位于第9染色体长臂区间RM6570-RM5652的qGY9在非条件及全生育期、株高、穗数、粒数和百粒重调整到同一水平后均可检测到,但加性效应、贡献率并不相同,显示该区间来自C堡的片段能够增加株高、穗数和百粒重从而增加产量。通过条件方法在第3染色体长臂区间RM7097-RM448及第6染色体长臂区间RM162-RM5753上定位到的产量QTL增加籽粒产量的等位基因可以降低株高,缩短生育期。     

水稻顶部三叶与穗重的关系及其QTL分析

 摘要: 对水稻汕优63重组自交系群体顶部3张叶片的长、宽、重和单穗重等10个性状进行了相关分析和QTL定位。穗重与9个叶片性状存在极显著的正相关，其中与倒2叶重的相关系数最大，剑叶重次之。所有性状在重组自交系群体中均存在双向超亲分离，接近正态分布。共检测到44个主效QTL和43对双位点互作影响上述10个性状。主效QTL分布于水稻的除第8染色体外的其余11条染色体上，贡献率介于3.19%~26.23%；互作分布于水稻的12条染色体上，贡献率变幅为2.03%~8.93%。第2染色体的R2510-RM211标记区间同时检测到控制单穗重和倒2叶重的QTL，该QTL对超级稻株型育种具有应用价值。     

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.

     

Mapping QTL for Heat-Tolerance at Grain Filling Stage in Rice

A mapping population of 98 lines (backcross inbred lines, BILs) derived from a backcross of Nipponbare/Kasalath// Nipponbare was planted at two experimental sites, Nanjing and Nanchang, and treated with high and optimal temperature during grain filling, respectively. The grain weight heat susceptibility index [GWHSI= (grain weight at optimum temperature-grain weight at high temperature) / grain weight at optimum temperature ×100] was employed to evaluate the tolerance of rice to heat stress. A genetic linkage map with 245 RFLP markers and a mixed linear-model approach was used to detect quantitative trait loci (QTLs) and their main effects, epistatic interactions and QTL×environment interactions (Q×E). The threshold of LOD score=2.0 was used to detect the significance of association between marker and trait. A total of 3 QTLs controlling heat tolerance during grain filling were detected, on chromosomes 1, 4 and 7, with LOD scores of 8.16, 11.08 and 12.86, respectively, and they explained the phenotypic variance of 8.94, 17.25 and 13.50 %, correspondingly. The QTL located in the C1100-R1783 region of chromosome 4 showed no QTL×environment interaction and epistatic effect, suggesting that it could be stably expressed in different environments and genetic backgrounds, and thus it would be valuable in rice breeding for heat tolerance improvement. This QTL allele, derived from Kasalath reduced 3.31% of the grain weight loss under heat stress. One located between R1613-C970 on chromosome 1 and the other between C1226-R1440 on chromosome 7, with additive effect 2.38 and 2.92%, respectively. The tolerance alleles of both these QTLs were derived from Nipponbare. Both of these QTLs had significant QTL×environment interactions, and the latter was involved in epistatic interaction also. Eight pairs of epistatic effect QTLs were detected, one pair each on chromosomes 1,2,3, 5, 7, 8, 10 and 12. The results could be useful for elucidating the genetic mechanism of heat-tolerance and the development of new rice varieties with heat tolerance during grain filling phase.     

水稻生理特性与抗旱性的相关分析及QTL定位

利用籼稻品种IR64和粳稻品种Azucena杂交产生的包含110个加倍单倍体株系的群体,在干旱胁迫和正常水分条件下,连续在2004年和2005年于抽穗期分别测定了叶片水势、相对含水量、叶绿素含量(SPAD值)、游离脯氨酸含量、气孔导度和蒸腾速率,并于成熟期取样,计算抗旱系数。与正常水分状况下相比,干旱胁迫条件下叶片的游离脯氨酸含量的增加达极显著水平,干旱胁迫条件下叶片的相对含水量、水势、叶绿素含量和气孔导度的降低均达显著或极显著水平。相关分析表明,在干旱胁迫条件下,叶片相对含水量、叶片水势与抗旱系数呈显著或极显著正相关。 利用175个RFLP标记构建的遗传连锁图谱分析了与抗旱性相关的叶片生理指标,共检测到与抗旱性相关的6个生理指标的7个加性QTL,31对上位性QTL,其中有2个主效QTL、9对上位性QTL存在环境互作效应。在两种水分条件下检测到的QTL结果有较大差异,说明干旱胁迫对控制与抗旱性相关的叶片生理性状基因的表达有显著的影响。在6个抗旱相关生理指标中,检测到的控制叶片气孔导度和水势的QTL较多,有3个加性QTL和8对上位性QTL控制气孔导度,有8对上位性QTL控制水势。     

水稻低温发芽力的QTL定位

以珍汕97B与多年生稻种质AAV002863的DH群体(198个株系)构建了包含140个SSR标记的连锁图谱,检测了影响水稻低温发芽力性状的数量性状座位（QTL）。15℃下处理6 d,两亲本珍汕97B与多年生稻AAV002863的发芽率分别为79.7%和30.1%,DH群体间的发芽率变化在0%~100%。QTL定位分析检测到2个与低温发芽力相关的基因座,分别位于第3和第10染色体上,贡献率分别为12.6%和12.9%,增效等位基因分别来自多年生稻AAV002863和珍汕97B。上位性分析结果显示,第1与第10染色体上存在影响低温发芽力的互作位点,其互作可以提高低温发芽力,参与互作的第10染色体上的位点也具有加性主效应。     

QTL Mapping of Low Temperature on Germination rate of Rice

Rice is a low temperature-sensitive crop especially during seedling, tillering, panicle development and flowering stages, which can easily be injured by cold treatment. Most of the high-yielding and high quality varieties can not be used in direct sowing because of low germination rate at low temperature (LTG). The reduction in seedling growth of rice due to low temperature is one of the major problems in tropical and subtropical areas at high altitude as well as in areas where cold mountain …     

稻米粒形和垩白度的QTL定位和上位性分析

 利用由181个家系组成的Lemont/特青籼粳交重组自交系群体,以及由161个RFLP、SSR标记和3个形态标记构建的全长为1916.5 cM、覆盖水稻基因组12 条染色体的连锁图,采用线性模型的复合区间作图方法（QTLMapper V10）,对粒长、粒宽、长宽比和垩白度等4个稻米品质性状的数量性状座位（QTL）进行了分析。在水稻的所有12 条染色体上共定位到7个加性主效QTL和19对上位性QTL,其中控制粒长、粒宽、长宽比的主效QTL各2个,控制垩白度的QTL 1个,分别解释12.8%、40.0%、26.0%和42.1%的表型变异;共检测到6对影响垩白度、6对影响粒长、7对影响长宽比的上位性QTL,分别解释52.2%、31.3%和38.2% 的表型变异。结果表明,上位性QTL和主效QTL一样在稻米粒形和垩白度的遗传中起着重要的作用。     

应用分子标记检测水稻耐盐性的QTL

 利用特三矮2号/CB组合构建了重组自交系群体(RI)。以60个RFLP标记检测142个纯系的基因型。在含有NaCl的电导率为12 dS/m的培养液中鉴定这些纯系的耐盐性。结果表明,RI群体的耐盐性出现超亲分离。构建了一张覆盖11条染色体、含52个标记位点的连锁图。仅检测到一个位于第5染色体的位点(RG13)显著与耐盐性有连锁。该位点的表型贡献率为11.6%。来自母本的该位点可提高耐盐性。分别对RG13与其它59个标记位点间的互作做检测,仅发现3对互作显著,即RG13×RG104; RG13×RG143; RG13×RG716。当来自母本的RG13分别与来自父本的RG104和RG143重组时,均明显提高耐盐性。分别来自母本的RG13和RG716能产生提高耐盐性的互作。这些基因互作结果为超亲分离提供了理论依据。     

QTL Analysis for Traits Associated with Feeding Value of Straw in Rice(Oryza sativa L.)

Genetic improvement of the digestibility of rice strew for increasing the utilization of the whole rice plant as feedstuffs is an important way to solve the feedstuffs shortage in southeastem China. To elucidate the genetic basis of the traits affecting the digestibility of rice straw, a rice population of 111 recombinant inbred lines (RILs) derived from a cross of Nekken 2xGaya was used to map the quantitative traits loci (QTLs) for in vitro dry matter digestion (IVDMD), the percentages of nonstructurel carbohydrate (NSC) and acid detergent fiber (exclusive of residual ash, ADFom) in 2005 and 2006. IVDMD was positively correlated with NSC, and negatively correlated with ADFom. A total of 16 QTLs were detected in the two years, and the amounts of variation explained by individual QTLs ranged from 6.9% to 15.5%. Some QTLs for IVDMD and ADFom on chromosome 2 were detected in the two years. On chromosome 2, the interval RM475-RM3515 contained QTLs for IVDMD in the two years and for NSC in 2006 only, while the interval RM3874-RM5305 influenced ADFom in the two years and NSC in 2005 only. At those loci, Gaya contributed favorable alleles to IVDMD and NSC, while Nekken 2 contributed positive alleles to ADFom.     

Molecular mapping of QTLs for gluten strength as measured by sedimentation volume and mixograph in durum wheat (Triticum turgidum L. ssp durum)

Quantitative trait loci (QTLs) responsible for gluten strength measured by SDS-sedimentation volume (SV), mixograph and grain protein content (GPC) were located on the molecular linkage map of a durum wheat recombinant inbred line population. QSv.macs-1B.1 flanked by Xgwm550–Glu-B3 was the most consistent QTL for SV identified in all the environments. The same QTL was also associated with mixograph peak energy, peak time and total energy. The Glu-B1 locus was at the center of another QTL responsible for SV, while, Glu-B2 influenced SV as well as mixograph peak energy and peak time. Apart from glutenin coding loci, QTLs influencing mixing parameters and GPC were located in three other marker intervals Xwmc48.2–Xpsp3030 (4B), Xgwm573–Xbarc231.1 (7A) and Xgwm46–Xgwm540.1 (7B). A total of 26 main effect QTLs and 10 digenic epistatic interactions (QQ) for quality traits were distributed over 11 chromosomes. Out of these, seven main effect QTLs and three QQ interactions were involved in interactions with environments (QE, QQE). The results indicated that along with chromosome 1B, chromosomes 4B, 7A and 7B are also important for improvement of gluten strength and GPC in durum wheat.     

Analysis of QTLs for Flag Leaf Shape and Its Response to Elevated CO2 in Rice （Oryza sativa）

To understand the responses of flag leaf shape in rice to elevated CO2 environment and their genetic characteristics,quantitative trait loci(QTLs)for flag leaf shape in rice were mapped onto the molecular marker linkage map of chromosome segment substitution lines(CSSLs)derived from a cross between a japonica variety Asominori and an indica variety IR24 under free air carbon dioxide enrichment(FACE,200μmol/mol above current levels)and current CO2 concentration(Ambient,about 370μmol/mol).Three flag-leaf traits,flag-leaf length(LL),width(LW)and the ratio of LL to LW (RLW),were estimated for each CSSL and their parental varieties.The differences in LL,LW and RLW between parents and in LL and LW within IR24 between FACE and Ambient were significant at 1%level.The continuous distributions and transgressive segregations of LL,LW and RLW were also observed in CSSL population,showing that the three traits were quantitatively inherited under both FACE and Ambient.A total of 16 QTLs for the three traits were detected on chromosomes 1,2,3,4,6,8 and 11 with LOD(Log10-likelihood ratio)scores ranging from 3.0 to 6.7.Among them,four QTLs (qLL-6*,qLL-8*,qLW-4*,and qRLW-6*)were commonly detected under both FACE and Ambient.Therefore,based on the different responses to elevated CO2 in comparison with current CO2 level,it can be suggested that the expressions of several QTLs associated with flag-leaf shape in rice could be induced by the high CO2 level.     

QTL Mapping for Grain Size Traits Based on Extra-Large Grain Rice Line TD70

Grain size traits, including grain length, grain width and grain thickness, are controlled by quantitative trait loci （QTLs）. Many QTLs relating to rice grain size traits had been reported, but their control mechanisms have not yet been elucidated. A recombinant inbred line （RIL） population of 240 lines, deriving from a cross between TD70, an extra-large grain size japonica line with 80 g of 1000-grain weight, and Kasalath, a small grain size indica variety, were constructed and used to map grain size QTLs to a linkage map by using 141 SSR markers in 2010 and 2011. Five QTLs for grain length, six for grain width and seven for grain thickness were detected distributing over chromosomes 2, 3, 5, 7, 9 and 12. Seven QTLs, namely qGL3.1, qGW2, qGW2.2, qGW5.1, qGW5.2, qGT2.3 and qGT3.1, were detected in either of the two years and explained for 56.19%, 4.42%, 29.41%, 10.37%, 7.61%, 21.19% and 17.06% of the observed phenotypic variances on average, respectively. The marker interval RM1347-RM5699 on chromosome 2 was found common for grain length, grain width and grain thickness; qGL3.1 and qGT3.1 were mapped to the same interval RM6080-RM6832 on chromosome 3. All 18 QTL alleles were derived from the large grain parent TD70. Most of the QTLs mapped in the present study were found the same as the genes previously cloned （GW2, GS3 or qGL3, GW5 and GS5）, and several were the same as the QTLs （GS7 and qGL-7） previously mapped. Three QTLs, qGL2.2 on chromosome 2, qGW9 and qGT9 on chromosome 9, were first detected. These results laid a foundation for further fine mapping or cloning of these QTLs.     

Mapping QTLs for Drought Tolerance at Seedling Stage in Rice Using Doubled Haploid Population

QTLs for drought tolerance at the rice seedling stage were analyzed using a doubled haploid (DH) population consisted of 251 lines from the cross between a japonica parent Maybelle and an indica parent Baiyeqiu. A genetic linkage map with 226 SSR marker loci was constructed. Single-locus analysis following composite interval mapping (CIM) detected a total of five QTLs located on five different chromosomes of rice. Four QTLs were also detected following two-locus analysis, resolving two pairs of epistatic QT...     

Dissecting Genetic Basis of Deep Rooting in Dongxiang Wild Rice

Deep rooting is an important trait in rice drought resistance. Genetic resources of deep-rooting varieties are valuable in breeding of water-saving and drought-resistant rice. In the present study, 234 BC₂F₇ backcross introgression lines were derived from a cross of Dongye 80 (an accession of Dongxiang wild rice as the donor parent) and R974 (an indica restorer line as the recurrent parent). A genetic linkage map containing 1 977 bin markers was constructed by ddRADSeq for QTL analysis. Thirty-one QTLs for four root traits (the number of deep roots, the number of shallow roots, the total number of deep roots and the ratio of deep roots) were assessed on six rice chromosomes in two environments (2020 Shanghai and 2021 Hainan). Two of the QTLs, qDR5.1 and qTR5.2, were located on chromosome 5 in a 70-kb interval. They were detected in both environments. qDR5.1 explained 13.35% of the phenotypic variance in 2020 Shanghai and 12.01% of the phenotypic variance in 2021 Hainan. qTR5.2 accounted for 10.88% and 10.93% of the phenotypic variance, respectively. One QTL (qRDR2.2) for the ratio of deep roots was detected on chromosome 2 in a 210-kb interval and accounted for 6.72% of the phenotypic variance in 2020. The positive effects of these three QTLs were all from Dongxiang wild rice. Furthermore, nine and four putative candidate genes were identified in qRDR2.2 and qDR5.1/qTR5.2, respectively. These findings added to our knowledge of the genetic control of root traits in rice. In addition, this study will facilitate the future isolation of candidate genes of the deep-rooting trait and the utilization of Dongxiang wild rice in the improvement of rice drought resistance.     

QTL mapping rolling,stomatal conductance and dimension traits of excised leaves in the Bala × Azucena recombinant inbred population of rice

The identification of markers linked to genes contributing to drought resistance promises opportunities to breed high yielding rice varieties for drought prone areas. Several studies using different mapping populations have previously identified quantitative trait loci (QTLs) for traits theoretically related to drought resistance. A mapping population of 176 F₆ recombinant inbred lines (RILs) derived from two upland rice varieties with contrasting aboveground drought avoidance traits (Bala and Azucena) with a linkage map of 157 markers was used to map QTLs for aboveground leaf morphological and physiological traits related to drought avoidance. Plants were grown for 6 weeks under controlled environmental conditions with three replications. Leaves were excised and placed on a balance. The rate of leaf rolling and water loss was recorded, after which leaf area, dry weight and specific leaf area were characterized. A simple method of estimating time to stomatal closure was employed. A total of 13 QTLs were detected for leaf morphological traits, three for initial transpiration and four for the proportion of water loss required to reach a specific advanced state of leaf rolling. No QTLs were detected for time of stomatal closure or speed of leaf rolling, nor for either water loss or transpiration at stomatal closure despite clear parental differences and moderate heritabilities in most of these traits. The co-location of QTLs for traits measured here and for drought avoidance previously reported from field experiments on chromosome 1, 3 and 5 link the genetics of drought resistance to leaf dimensions and physiology. However, a physiological explanation for a QTL for drought avoidance on chromosome 7 remains elusive.     

Marker-assisted genetic analysis of non-acclimated freezing tolerance and cold acclimation capacity in a backcrossSolarium population

Random amplified polymorphic DNA (RAPD) and simple sequence repeat (SSR) markers were used to construct a partial genetic linkage map in a potato backcross population. The population, derived from two diploid wildSolatium species (frost tolerant, able to cold acclimateS. commersonii; frost sensitive, unable to cold acclimateS. cardiophyllum), was used to map quantitative trait loci (QTL) of non-acclimated relative freezing tolerance (NARFT) and cold acclimation capacity (CAC). Precise assessment of these traits allowed distinction of small but significant differences among 35 backcross genotypes. NARFT and CAC were not correlated in the segregating population, suggesting independent genetic control for these two major components of freezing tolerance. The linkage map spanned 479.4 cM and included 77 RAPD markers and two SSR markers, with 38 RAPD and 10 SSR unassigned markers. Two QTLs for NARFT were detected in two different linkage groups, accounting for 44.0% of the phenotypic variation for this trait. Two QTLs for CAC were detected, accounting for 24.9% of the phenotypic variation for this trait. QTLs for NARFT and CAC were detected at separate genomic regions, in support of the independent genetic control of these two traits. QTLs for NARFT and CAC were detected in a linkage group identified as part of chromosome V, suggesting that such chromosome constitutes a prime candidate for fine-mapping. Due to the relatively small progeny size evaluated in this study, additional QTLs for NARFT and CAC could have been involved but not identified. Therefore, the conclusions derived from this study should be considered preliminary.