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.     

利用单片段代换系研究水稻产量相关性状QTL加性及上位性效应

产量及其相关性状如单株有效穗数、千粒重、穗实粒数、穗总粒数和结实率等是水稻重要的农艺性状,了解产量及其相关性状QTL的加性及上位性效应对以分子标记聚合育种改良水稻产量具有重要意义。本文以16个单片段代换系及15个双片段代换系分析了水稻产量相关性状QTL的加性及上位性效应。共检出影响产量及其相关性状的13个QTL,包括产量性状1个、单株有效穗数1个、千粒重4个、穗实粒数4个、穗总粒数2个和结实率1个,分布于第2、第3、第4、第7和第10染色体上。此外,检出12对双基因互作。结果显示,2个正向(或负向)产量性状QTL聚合,往往会产生负向(或正向)的上位性效应,能否产生更大(或更小)的目标性状,取决于双片段遗传效应(加性效应与上位效应代数和)绝对值与单片段最大加性效应绝对值的差。本研究结果对实施高产分子标记聚合育种方法有重要参考价值。     

不同生态环境下玉米产量性状QTL分析

以玉米（Zea mays L.）自交系黄早四和Mo17为亲本得到的191个F₂单株为作图群体,衍生的184个F_2∶3 家系作为性状评价群体,分析了单株穗数、穗行数、行粒数、百粒重和单株籽粒产量在北京和新疆2个生态环境下的表现和数量性状基因位点的定位结果。QTL检测结果表明,2个环境共检测出47个QTL,分布于除第10染色体以外的9条染色体,其中与单株穗数相关的QTL共10个,可解释的表型变异为5.3%~25.6%;与穗行数相关的QTL共13个,可解释的表型变异为4.5%~23.2%;与行粒数相关的QTL有9个,解释的表型变异为5.4%~13.7%;与百粒重相关的QTL达10个,可解释的表型变异为4.9%~13.3%;与单株籽粒产量相关的QTL有5个,可解释的表型变异为6.1%~35.8 %。大部分产量QTL只在单一环境下被检测到,说明产量相关QTL与环境之间存在明显的互作。表型相关显著的产量性状,它们的QTL容易在相同或相邻标记区间检测到。研究还发现了若干个QTL富集区域,可能是发掘通用QTL的候选位点。     

Identification of a novel QTL for the number of spikelets per panicle using a cross between indica‐ and japonica‐type high‐yielding rice cultivars in Japan

Yield is a complex trait. To improve it, the accumulation of the favourable alleles of valuable genes is required for each yield‐related trait. In this study, we used two high‐yielding rice cultivars developed in Japan, indica‐type ‘Takanari’ and japonica‐type ‘Momiroman’, for a genetic analysis of the sink capacity‐related traits. An F₂ population showed transgressive segregation for the number of spikelets per panicle. Quantitative trait locus (QTL) analysis detected four QTLs for the trait. Two of the QTLs were most likely identical to previously cloned GN1a and APO1, and their Takanari alleles had positive effects. The Momiroman alleles of the other two QTLs had positive effects, and one of these QTLs was most likely identical to SPIKE/GPS. The QTL on the long arm of chromosome 3 appeared to be novel; it clustered with QTLs for grain length and days‐to‐heading. Substitution mapping revealed that the close linkage of QTLs caused the clustering. These results suggest that the combination of the favourable alleles of detected QTLs could lead to greater sink capacity than that of the parental cultivars.     

Mapping quantitative trait loci (QTLs) underlying seed vitamin E content in soybean with main,epistatic and QTL × environment effects

Soybean (Glycine max (L.) Merr.) seed contains small amounts of tocopherol, a non‐enzymatic antioxidant known as lipid‐soluble vitamin E (VE). Dietary VE contributes to a decreased risk of chronic diseases in humans and has several beneficial effects on resistance to stress in plants, and increasing VE content is an important breeding goal for increasing the nutritional value of soybean. In this study, quantitative trait loci (QTLs) underlying VE content with main, epistatic and QTL × environment effects were identified in a population of F_5 : 6 recombinant inbred lines from a cross between ‘Hefeng 25’ (a low‐VE cultivar) and ‘OAC Bayfield’ (a high‐VE cultivar). A total of 18 QTLs were detected that showed additive main effects (a) and/or additive × environment interaction effects (ae) in different environments. Moreover, 19 epistatic pairs of QTLs were found to be associated with α‐tocopherol (α‐Toc), γ‐tocopherol (γ‐Toc), δ‐tocopherol (δ‐Toc) and total VE (TE) contents. The QTLs identified in multienvironments could provide more information about QTL by environment interactions and could be useful for the marker‐assistant selection of soybean cultivars with high seed VE contents.     

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.     

小麦籽粒产量及穗部相关性状的QTL定位

由小麦品种花培3号和豫麦57杂交获得DH群体168个株系,种植于3个环境中,利用305个SSR标记对籽粒产量和穗部相关性状(穗长、穗粒数、总小穗数、可育小穗数、小穗着生密度、千粒重和粒径)进行了QTL定位。利用基于混合线性模型的QTLNetwork 2.0软件,共检测到27个加性效应和13对上位效应位点,其中 8个加性效应位点具有环境互作效应。相关性高的性状间有一些共同的QTL位点,表现出一因多效或紧密连锁效应。5D染色体区段Xwmc215–Xgdm63,检测到控制籽粒产量、穗粒数、总小穗数、可育小穗数和小穗着生密度5个性状的QTL位点,各位点的遗传贡献率较大且遗传效应方向相同,增效等位基因均来源于豫麦57,适用于分子标记辅助育种和聚合育种。控制千粒重与穗粒数的QTL位于染色体不同区段,有利于实现穗粒数与粒重的遗传重组。     

优良水稻染色体片段代换系Z746的鉴定及重要农艺性状QTL定位及其验证

粒型、株高及穗部组成性状与产量形成密切相关,是水稻重要农艺性状,但遗传基础复杂。染色体片段代换系是用于复杂性状遗传研究的良好材料。本研究鉴定了一个以日本晴为受体、西恢18为供体亲本的水稻优良染色体片段代换系Z746。Z746携带来自西恢18的7个代换片段,平均代换长度为3.99 Mb,其株高、粒长和穗部性状均与受体存在显著差异。进一步通过日本晴与Z746杂交构建的次级F2群体共检测到36个相关QTL,分布于2号、3号、4号、6号和11号染色体。其中5个可能与已克隆基因等位,如qPH3-1等, 8个可被多次检出,表明这些是遗传稳定的主效QTL。Z746的粒长主要由4个QTL(qGL3、qGL4、qGL2、qGL6)控制,其中qGL3和qGL4对粒长变异的贡献率分别为60.28%和27.47%。株高由5个QTL控制,穗长由4个QTL控制,每穗粒数由2个QTL控制,千粒重由2个QTL控制。然后以MAS在F3共选出8个单片段代换系,并以此在F4进行了相关QTL验证,共有24个QTL可被8个单片段代换系(SSSL)检出,重复检出率为66.7%,表明这些QTL遗传稳定。本研究为目的 QTL的进一步...     

Mapping quantitative trait loci influencing panicle‐related traits from Chinese common wild rice (Oryza rufipogon) using introgression lines

Panicle‐related traits are important agronomic traits which directly associated with grain yield. In this study, we investigated quantitative trait loci (QTLs) associated with panicle‐related traits using a set of 265 introgression lines (ILs) of common wild rice (Oryza rufipogon Griff.) in the background of Indica high‐yielding cultivar Guichao 2 (O. sativa L.). A total of 39 QTLs associated with panicle‐related traits including panicle length (PL), primary branch number (PBN), secondary branch number (SBN), spikelet number per panicle (SPP) and spikelet density (SD), were detected in the ILs with single‐point analysis. The alleles of 20 QTLs derived from wild rice showed positive effects, and some QTLs, such as, QPl1b for PL, QPbn8 for PBN, QSd4 and QSd11b for SD and QSpp4 for SPP showed larger positive effects, providing good candidates and useful information for marker‐aided improvement of yield potential of rice. Most of the QTLs controlling SPP, SBN and SD were located in cluster or closely linked on chromosomes, and the directions of their additive effects were consistent, which explained the genetic basis of significant correlations between their phenotypic characters.     

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。     

Fine mapping of a QTL for the number of spikelets per panicle by using near‐isogenic lines derived from an interspecific cross between Oryza sativa and Oryza minuta

We constructed a high‐resolution physical map for the qSPP7 QTL for spikelets per panicle (SPP) on rice chromosome 7 across a 28.6‐kb region containing four predicted genes. Using a series of BC₇F₄ near‐isogenic lines (NILs) derived from a cross between the Korean japonica cultivar ‘Hwaseongbyeo’ and Oryza minuta (IRGC Acc. No. 101144), three QTLs for the number of SPP, grains per panicle and primary branches were identified in the cluster (P ≤ 0.01). All three QTLs were additive, and alleles from the O. minuta parent were beneficial in the ‘Hwaseongbyeo’ background. qSPP7 was mapped to a 28.6‐kb region between the two simple sequence repeat (SSR) markers RM4952 and RM21605. The additive effect of the O. minuta allele at qSPP7 was 23 SPP, and 43.6% of the phenotypic variance was explained by the segregation of the SSR marker RM4952. Colocalization of the three QTLs suggested that this locus was associated with panicle structure and had pleiotropic effects. The NIL populations and molecular markers are useful for cloning qspp7.     

Construction of chromosomal segment substitution lines and genetic dissection of introgressed segments associated with yield determination in the parents of a super‐hybrid rice

Heterosis is a phenomenon whereby hybrids of inbred lines produce favourable phenotypes that exceed those of their parents. Traits of interest are higher yield and stronger stress tolerance. The two‐line super‐hybrid rice ‘Liangyoupei9’ (LYP9) shows superiority to both its elite inbred line ‘93‐11’ and ‘Pei'ai64s’ (‘PA64s’) parents and conventional hybrids. However, the genetic basis of its hybrid vigour, especially yield determination, remains elusive. In the present study, a set of 156 chromosome segment substitution lines (CSSLs) carrying overlapping segments from ‘PA64s’ in a genetic background of ‘93‐11’ were constructed and planted in six environments. Three major agronomic traits, viz. panicle length (PL), heading date (HD) and plant height (PH), and five yield‐related traits, viz. grain weight per panicle (GWP), number of grains per panicle (GPP), 1000‐grain weight (TGW), seed set (SS) and number of panicles of per plant (PPP), were evaluated over 3 years. Quantitative trait loci (QTL) analysis was conducted using a likelihood ratio test based on stepwise regression. Forty‐six putative QTL distributed on 11 chromosomes were detected in more than one year. Remarkably, GWP of four CSSLs carrying positive yield QTL outperformed the recurrent parent ‘93‐11’ by more than 15%, in at least two environments. These results indicate that CSSLs are effective in identifying yield‐associated traits, and lines harbouring such QTL will be rich in resources for future molecular breeding programmes.     

Detection of epistatic and environmental interaction QTLs for leaf orientation‐related traits in maize

Leaf architecture traits in maize are quantitative and have been studied by quantitative trait loci (QTLs) mapping. However, additional QTLs for these traits require mapping and the interactions between mapped QTLs require studying because of the complicated genetic nature of these traits. To detect common QTLs and to find new ones, we investigated the maize traits of leaf angle, leaf flagging‐point length, leaf length and leaf orientation value using a set of recombinant inbred line populations and single nucleotide polymorphism markers. In total, 19 QTLs contributed 4.13–13.52% of the phenotypic effects to the corresponding traits that were mapped, and their candidate genes are provided. Common and major QTLs have also been detected. All of the QTLs showed significant additive effects and non‐significant additive × environment effects in combined environments. The majority showed additive × additive epistasis effects and non‐significant QTL × environment effects under single environments. Common and major QTLs provided information for fine mapping and gene cloning, and SNP markers can be used for marker‐assisted selection breeding.     

影响水稻穗部性状及籽粒碾磨品质的QTL及其环境互作分析

利用优质恢复系测258为轮回亲本与粳型糯稻新品系IR75862杂交创制的BC₁F₇回交导入系群体,在广西南宁和海南三亚定位了产量相关性状(二次枝梗数、穗总粒数、穗实粒数、粒重和穗重)、粒型(粒长、宽、厚)和碾磨品质(糙米率、精米率和整精米率)的主效QTL并剖析其环境互作效应。双亲在穗实粒数、千粒重、粒长和粒宽及整精米率等性状上存在显著差异。各产量相关性状间呈极显著正相关,而与千粒重和粒长呈极显著负相关。多数产量及粒型相关性状与3种碾磨品质相关不显著。在南宁和三亚环境下检测到影响产量相关性状、粒型及碾磨品质的主效QTL共计57个,包括二次枝梗数的6个,穗实粒数4个,穗总粒数、粒重和穗重各5个,粒长9个,粒宽7个,粒厚1个,糙米率4个,精米率5个和整精米率6个,分布在除第11染色体外的所有染色体上。多数影响枝梗数、穗粒数和粒重的QTL成簇分布,而且与影响BR、MR和HR的QTL分布在不同染色体区域。在第2、第3、第4、第5和第6染色体上鉴定出影响穗粒数、粒重、粒型及碾磨品质的重要QTL,这些QTL在以往不同遗传背景和环境下被多次检测到。在第8染色体RM152~RM310区间鉴定到1个影响粒长和粒宽的新的QTL,能同步增加粒宽和粒长。鉴定出的这些稳定表达的QTL具有标记辅助选择育种的应用价值。整精米率是受环境影响最大的性状,其QTL的环境互作效应明显。对QTL的环境互作效应特点及其在品种标记辅助改良中的作用进行了深入探讨。     

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.     

Identification and validation of number of pod ‐ and seed ‐ related traits QTLs in soybean

Traits related to the number of pods and seeds are important yield factors on soybean. The relationships between phenotype and quantitative trait loci (QTLs) of these traits may reveal the mechanisms underlying productivity. Our study objectives were to analyse phenotypic correlations, detect stable QTLs and identify candidate genes useful for marker‐assisted selection. Phenotypic analyses revealed that NThSP (number of three‐seeded pods) was positively correlated with NPPP (number of pods per plant) and SNPP (number of seeds per plant). Seventy‐five QTLs were identified based on the mean phenotypic data for at least 2 years. We detected two to 15 and one to three significant QTLs identified at the same location, respectively. Six consensus QTLs associated with at least two NPS‐related (number of pods and seeds related) traits were identified. Two of these were verified in another population. The QTLs for NPPP, SNPP and NThSP formed a consensus QTL cluster on GM02. Another 27 QTLs also formed clusters in five regions. Fifteen candidate genes were mined and discussed. The results will provide more information to soybean breeding.     

水稻千粒重和垩白粒率的QTL及其互作分析

产量因子千粒重和稻米品质指标垩白粒率密切相关。本研究以越光/Kasalath//越光BIL群体为材料,分析千粒重和垩白粒率的相关性、QTL、上位性互作及其环境的互作效应。相关分析表明,群体千粒重和垩白粒率在2005年和2006年均呈极显著正相关,相关系数分别为0.42和0.35 (P<0.001)。2年共检测到千粒重QTL 11个,其中5个在2年重复检测到,5个具有环境互作效应;千粒重上位性互作8对,7对与环境存在互作。垩白粒率QTL 6个,3个具有环境互作效应;上位性互作9对,其中4对具有上位性环境互作效应。比较分析发现3个主效QTL同时控制千粒重和垩白粒率的表现,千粒重和垩白粒率的增效等位基因来自同一亲本;1对上位性互作同时对千粒重和垩白粒率有相同的影响。一些与垩白粒率不相关的千粒重主效QTL,如qTGW-3c、qTGW-4a和qTGW-6b,可为育种所利用。对利用QTL定位结果进行千粒重和垩白粒率分子辅助选择育种进行了探讨。     

利用东乡普通野生稻染色体片段置换系定位产量相关性状QTL

以东乡普通野生稻和日本晴为亲本构建的染色体片段置换系为研究材料, 2019年分别在北京、山东临沂和江西南昌对分蘖数、穗粒数和粒形等11个产量相关性状进行多环境鉴定,结合染色体片段置换系基因型数据定位水稻产量相关性状QTL。3个环境共检测到68个QTL,包括株高4个、穗长5个、分蘖数2个、一次枝梗数7个、一次枝梗粒数8个、二次枝梗数8个、二次枝梗粒数10个、每穗粒数6个、千粒重7个、粒长8个和粒宽3个; LOD值介于2.50～12.66之间,贡献率变幅为4.67%～27.79%,15个QTL的贡献率大于15%;24个QTL与已报道位点/基因位置重叠,44个QTL为新发现位点; 6个QTL在2个环境能被检测到, 1个QTL qTGW2能在3个环境检测到,且是还未报道的新位点。最后,利用BSA法验证了qPH7、qPBPP8-2和qGW10三个QTL的可靠性。本研究将为后续产量相关性状基因克隆以及进一步解析其遗传基础和分子调控机制奠定基础。     

用单片段代换系（SSSLs）研究水稻株高及其构成因素QTL加性及上位性效应

株高是典型的数量性状,易受遗传背景和环境等因素的影响。单片段代换系和双片段聚合系减少了个体间遗传背景的干扰,是鉴定QTL和研究QTL上位性的新型遗传材料。本研究采用随机区组试验设计方法以初级单片段代换系间杂交衍生的16个次级单片段代换系和15个双片段聚合系分析了株高及其构成因素QTL的加性效应及加性×加性上位性效应。共鉴定出11个QTL,其中3个株高QTL,1个倒1节间长QTL,2个倒2节间长QTL,2个倒3节间长QTL和3个倒4节间长QTL,分布于第4、6和10染色体上。鉴定出23对双基因互作,其中7对为没有显著效应的座位间互作,16对为有显著效应的QTL与没有显著效应的座位间互作。结果表明,QTL加性效应和QTL间的上位性效应都是株高及构成因素的重要遗传组成。通过单片段代换系杂交衍生的次级单片段代换系和双片段聚合系可提高QTL鉴定和上位性分析的灵敏度。     

低磷胁迫下水稻产量性状变化及其QTL定位

为研究低磷胁迫对水稻产量及其构成因素QTL表达的影响, 以耐低磷旱稻IRAT109和磷敏感水稻越富杂交的116个株系的DH群体为材料, 在低磷和正常栽培条件下, 调查了千粒重、结实率、有效穗数、穗粒数及单株产量等性状。结果表明, 结实率、有效穗数和单株产量对低磷胁迫的敏感性较大, 而千粒重、穗粒数对低磷胁迫的敏感性较小。利用水稻分子连锁图中94个RFLP标记和71个SSR标记, 依据以上5性状低磷胁迫下与对照的差值进行QTL定位。共检测出17个QTL, 其中12个对表型变异的贡献率大于10%。3、6和7号染色体上3个标记区域存在QTL成簇分布, 这些高贡献率QTL及成簇分布QTL可作为水稻耐低磷产量性状分子育种的重要候选区域。