Identification of population-specific QTLs for flowering in soybean

Flowering is an important stage in plant development and crucial for adaptation of plant species to different environments. Two soybean mapping populations were used to identify quantitative trait loci (QTLs) for days to flowering (DF) and days to maturity (DM) by genotyping simple sequence repeat (SSR) markers. Single-factor analysis of variance detected association of phenotypic data with SSR markers in each population. DF QTLs were identified on four chromosomes (chrs.); two QTLs located on chrs. 2 and 13 with Satt041 and Satt206 in the Jinpumkong 2 × SS2-2 population and other two DF QTLs were detected on chrs. 6 and 19 with Satt100 and Satt373 in the Iksannamulkong × SS2-2 population. The major QTLs associated with Satt100 explained 30.3% of maximum phenotypic variation. Especially, all DF QTLs included QTLs for DM, except Satt206 on chr. 13. Moreover, two additional DM QTLs were mapped on chrs. 10 and 11 with Satt243 and Satt359, respectively. DF QTL on chr. 2 with Satt041 was the newly identified QTL only in the Jinpumkong 2 × SS2-2 population and explained 10.3% of the phenotypic variation. The single locus of Satt100 on chr. 6 and Satt373 on chr. 19 were located on soybean genomic regions of the known flowering gene loci E1 and E3, respectively. These population-specific QTLs (Satt100 and Satt373) are the major QTLs for flowering time, putatively, they may be related to maturity QTLs with large effect. Additionally, these QTLs are valuable for marker-assisted approaches and could be widely adopted by soybean breeders.     

QTL identification of yield-related traits and their association with flowering and maturity in soybean

Two soybean recombinant inbred line populations, Jinpumkong 2 × SS2-2 (J × S) and Iksannamulkong × SS2-2 (I x S) showed population-specific quantitative trait loci (QTLs) for days to flowering (DF) and days to maturity (DM) and these were closely correlated within population. In the present study, we identified QTLs for six yield-related traits with simple sequence repeat markers, and biological correlations between flowering traits and yield-related traits. The yield-related traits included plant height (PH), node numbers of main stem (NNMS), pod numbers per plant (PNPP), seed numbers per pod (SNPP), 100-seed weight (SW), and seed yield per plant (SYPP). Eighteen QTLs for six yield-related traits were detected on nine chromosomes (Chrs), containing four QTLs for PH, two for NNMS, two for PNPP, three for SNPP, five for SW, and two for SYPP. Two highly significant QTLs for PH and NNMS were identified on Chr 6 (LG C2) in both populations where the major flowering gene, E1, and two DF and DM QTLs were located. One other PNPP QTL was also located on this region, explaining 12.9% of phenotypic variation. Other QTLs for yield-related traits showed population-specificity. Two significant SYPP QTLs potentially related with QTLs for SNPP and PNPP were found on the same loci of Chrs 8 (Satt390) and 10 (Sat_108). Also, highly significant positive phenotypic correlations (P < 0.01) were found between DF with PH, NNMS, PNPP, and SYPP in both populations, while flowering was negatively correlated with SNPP and SW in the J × S (P < 0.05) and I × S (P < 0.01) populations. Similar results were also shown between DM and yield-related traits, except for one SW. These QTLs identified may be useful for marker-assisted selection by soybean breeders.     

Fine mapping of a major flowering time QTL on soybean chromosome 6 combining linkage and association analysis

Flowering time is a key trait in the plant life cycle and an important selection criterion for soybean. Here, we combine the advantages of genome-wide association and linkage mapping to identify and fine map quantitative trait loci (QTLs) associated with flowering time. Linkage mapping was performed using 152 recombinant inbred lines and a major QTL, qFT6, affecting flowering time was found on chromosome 6. To refine the qFT6, the 192 natural accessions were genotyped using eight new simple sequence repeats and 10 single nucleotide polymorphisms markers covering the qFT6 region Haplotype analysis showed that the haplotype between markers BARC-014947-01929 and Satt365 could explain more phenotypic variation (26.5 %) than any other combination of markers. These results suggested that the target flowering time gene was located in ~300 kb between BARC-014947-01929 and Satt365, including three predicted genes. High-resolution map in qFT6 region will be useful not only for marker-assisted selection of flowering time but also for further positional cloning of the target gene. These results indicate that combining association and linkage mapping provides an efficient approach for fine mapping of soybean genes.     

大豆苗期耐淹性的遗传与QTL分析

洪涝灾害是大豆生产的主要逆境之一,培育耐涝品种是抗灾保收的重要措施。大豆耐涝性育种方案的设计必须以耐涝性遗传为前提。以苏88-M21(淹水不敏感)×新沂小黑豆(淹水敏感)衍生的175个重组自交系(NJRISX)为材料,在盆栽V2期土壤表层保持5~7 cm水层20 d的淹水条件下,研究大豆苗期耐淹性的遗传和QTL定位。通过对8个耐淹性有关性状的相关分析和主成份分析,确定以处理前后株高变化量、处理终叶龄和成熟期株高3个性状的平均耐淹指数为评价指标。NJRISX家系间耐淹性差异极显著,存在超亲分离。主基因+多基因分离分析表明该群体的耐淹性为2对连锁主基因+多基因遗传,主基因遗传率为62.83%,多基因的遗传率为8.90%。WinQTLCart2.5复合区间及多区间QTL定位分析均检测到2个QTL,位于连锁群L2上的satt229~satt527和satt527~satt286区间,对表型的解释率分别为11.76%~25.20%和10.10%~12.34%。大豆NJRISX群体苗期耐淹性遗传分离分析与QTL定位结果相对一致。     

Analysis of quantitative trait loci underlying the period of reproductive growth stages in soybean (<Emphasis Type="Italic">Glycine max</Emphasis> [L.] Merr.)

Quantitative trait loci (QTLs) underlying reproductive growth stages are important for molecular breeding of soybeans [Glycine max (L.) Merr.]. Most of these QTLs identified so far derived from a single environment, and thus may be influenced by specific environmental conditions. In this study (from 2004 to 2005), analysis of QTLs underlying the period to reach a given reproductive growth stage was performed in three different environments (Harbin, Heilongjiang Province, China). QTL analysis was achieved with a recombination inbred line (RIL) population consisting of 153 lines. The RIL population derived from a cross between an American semi-dwarf cultivar (cv. Charleston) and a Chinese line with a short growth stage (cv. Dongnong 594). The growth stage data of soybean was recorded for each day. QTLs for all eight reproductive growth stages of soybean (R1 to R8) were analyzed by a composite interval mapping method combined with a mixed genetic model. Fifty-four QTLs displayed main effects and 56 QTL pairs showed epistatic effects. Two marker intervals (Satt173–Satt581, Satt402–Satt267), located on the linkage group O and D1a respectively, strongly influenced plant developmental processes during reproductive growth stages. The findings of this study open the possibility to modulate the structure of soybean growth stages by marker-assisted selection and pyramiding QTL analysis. H.-M. Qiu and D.-W. Xin contributed equally to this work.     

Quantitative trait loci for agronomic traits in soybean

There continues to be improvement in seed yields of soybean by conventional breeding, but molecular techniques may provide faster genetic gains. The objective of this study was to identify quantitative trait loci (QTL) associated with the agronomic traits seed yield, lodging, plant height, seed filling period and plant maturity in soybean. To achieve this objective, 101 F₆‐derived recombinant inbred lines (RIL) from a population developed from a cross of N87‐984‐16 × TN93‐99 were used. Experiments were conducted in six environments during 2002–2003. Heritability estimates on an entry mean basis from data combined across environments ranged from 0.12 to 0.65 for seed yield and seed filling period, respectively. Composite interval mapping detected one QTL for yield (near Satt076), two for lodging (near Satt225 and Satt593) and four for maturity (near Satt263, Satt292, Satt293 and Satt591) in this population. Additional environmentally sensitive QTL for these traits, and for seed filling period and plant height are also reported. The QTL associated with agronomic traits that we report and the recently released germplasm (PI 636460) from this population may be useful in soybean breeding programmes.     

Linkage fine-mapping and QTLs affecting morpho-agronomic traits of a Mesoamerican × Andean RIL common bean population

This paper proposes the construction of a genetic linkage map with 376 recombinant inbred lines (RILs) derived from a cross between Mesoamerican?×?Andean common bean (Phaseolus vulgaris L) parents based on single nucleotide polymorphism (SNP) markers; and to detect quantitative trait loci (QTLs) associated with seven morpho-agronomic traits: number of days to flowering (DF), number of days to maturity (DM) or crop cycle; plant architecture (ARC); seed yield (YLD); degree of seed flatness (SF); seed shape (SS); and 100-seed weight (SW). A total of 3060 polymorphic SNP markers were used and 2041 segregated at a 1:1 ratio in the RIL population, as expected. These markers were subjected to linkage analysis in each chromosome. The genetic linkage analysis resulted in linkage maps with a total of 1962 markers spanning 1079.21 cM. A total of 29 QTLs associated with seven morpho-agronomic traits were detected on the 11 chromosomes, which explained between 3.83 and 32.92% of the phenotypic variation in DF. A total of 18 candidate genes associated with the detected QTLs were identified and related with biological processes, molecular functions and cellular components.     

基于元分析的大豆生育期QTL的整合

共搜集整理了12年来已经报道的与大豆生育期有关的98个QTL,通过BioMercator2.1和公共标记映射整合到大豆公共遗传连锁图谱soymap2上,并利用元分析技术推断QTL位置,计算提取真正有效的QTL。发掘出大豆两个重要生育时期,共9个“真实QTL”及其连锁标记,其中与开花期(R1)相关的有7个,与成熟期(R8)相关的有2个,建立了QTL的一致性图谱,其中L连锁群上的一个定位区间包含一个已发表的有关R1的基因。在5个连锁群上共发现10个控制多个生育时期的QTL。本研究结果为大豆生育期QTL精细定位和基因克隆奠定了基础。     

Mapping quantitative trait loci for yield-related traits in soybean (Glycine max L.)

Development of soybean cultivars with high seed yield is a major focus in soybean breeding programs. This study was conducted to identify genetic loci associated with seed yield-related traits in soybean and also to clarify consistency of the detected QTLs with QTLs found by previous researchers. A population of 135 F_2:3 lines was developed from a cross between a vegetable soybean line (MJ0004-6) and a landrace cultivar from Myanmar ({"type":"entrez-nucleotide","attrs":{"text":"R18500","term_id":"772110","term_text":"R18500"}}R18500). They were evaluated in the experimental field of Kasetsart University, Kamphaeng Saen, Nakhon Pathom, Thailand in a randomized complete block design with two replications each in 2011 and 2012 growing seasons. The two parents exhibited contrasting characteristics for most of the traits that were mapped. Analysis of variance showed that the main effects of genotype and environment (year) were significant for all studied traits. Genotype by environment interaction was also highly significant for all the traits. The population was genotyped by 149 polymorphic SSR markers and the genetic map consisted of 129 SSR loci which converged into 38 linkage groups covering 1156 cM of soybean genome. There were 10 QTLs significantly associated with seed yield-related traits across two seasons with single QTLs explaining between 5.0% to 21.9% of the phenotypic variation. Three of these QTLs were detected in both years for days to flowering, days to maturity and 100 seed weight. Most of the detected QTLs in our research were consistent with earlier QTLs reported by previous researchers. However, four novel QTLs including SF1, SF2 and SF3 on linkage groups L and N for seed filling period and PN1 on linkage group D1b for pod number were identified in the present study.     

大豆耐旱选择群体QTL定位

以红丰11为轮回亲本、Clark为供体亲本构建回交群体进行耐旱性鉴定,对获得选择群体进行全基因组SSR标记扫描,计算供体基因型导入频率,利用卡方测验检测偏分离SSR位点,并结合GGT软件对各连锁群分析, 对5个耐旱相关性状进行QTL定位。以卡方测验检测到23个SSR偏分离位点(超导入),分布于10条连锁群。方差分析表明,8个叶片持水能力QTL分布于A₁、B₁、C₂、E、L和N连锁群;9个根长QTL分布于C₂、F、G和I连锁群;11个根干重QTL分布于A₂、B₁、B₂、E、F、K、L、M和O连锁群;12个产量QTL分布于B₁、D₁a、E、F、G、I、L、M和O连锁群;7个生物量QTL分布于E、F、G、K、L和N连锁群。在E连锁群的Sat_136位点,对于叶片持水能力、根干重、产量和生物量具有一致性;在F连锁群的GMRUBP位点,对于根干重和生物量具有一致性,Satt586位点,对于根长、根干重和产量具有一致性;在K连锁群的Satt167位点,对于根干重和生物量具有一致性,SOYPRP1位点,对于根长和生物量具有一致性;在L连锁群的Satt398位点,对于根长和产量具有一致性,Satt694位点对于叶片持水能力和生物量具有一致性;在M连锁群的GMSL514位点,对于根干重和产量具有一致性;以上位点均与卡方测验检测到的“超导入”位点具有一致性。经过供体等位基因卡方测验和耐旱QTL定位,共检测到33个QTL,其中有17个同时被检测到。这些位点可能是控制大豆耐旱性的重要位点。     

Identification of QTL underlying soluble pigment content in soybean stems related to resistance to soybean white mold (<Emphasis Type="Italic">Sclerotinia sclerotiorum</Emphasis>)

The frequency of soybean white mold (Sclerotinia sclerotiorum, SWM) outbreaks has increased since 1978, and this disease is currently considered to be the second most important cause of soybean yield loss worldwide. We have studied SWM in stems of soybean cultivar Maple Arrow, which shows partial resistance to SWM, in an attempt to identify the quantitative trait loci (QTLs) underlying soluble pigment(s) content, which is associated with SWM resistance. A SWM-susceptible cultivar, Hefeng 25, was crossed with Maple Arrow, and 149 F_5:6 recombinant inbred lines were subsequently advanced through single-seed-descent. A total of 109 simple sequence repeat (SSR) markers were used to construct a genetic linkage map. Three QTLs for soluble pigment content in soybean stems associated with the resistance to SWM, namely, Qsp-1 (Satt502-Sat_159), Qsp-2 (Sat_156-Satt251), and Qsp-3 (Satt525-Satt233), were identified in 2007 and 2008 and located onto linkage groups D1a+q, B1 and A2, respectively. The phenotypic variation (R ²) explained by these QTLs ranged from 6.29 to 15.37%. These three QTLs were not significantly related to known QTLs associated with escape resistant mechanisms. The use of these QTLs in marker-assisted selection may contribute to improved soybean resistance to SWM.     

大豆苗期耐淹性的遗传与QTL分析

涝害是世界上许多国家的重大自然灾害。耐涝性可分为耐湿(渍)性和耐淹性。以科丰1号(高度耐淹)×南农1138-2(不耐淹)衍生的RIL群体(NJRIKY)为材料, 以盆栽全淹条件下的存活率为耐淹性指标, 采用主基因+多基因混合遗传模型分离分析法进行遗传分析, 并利用WinQTL Cartographer Version 2.5程序的复合区间作图法(CIM)及多区间作图法(MIM)进行QTL定位。结果表明, 两次试验的耐淹性均存在超亲变异, 试验间、家系间以及试验与家系互作间的差异均极显著; NJRIKY大豆群体的耐淹性为3对等加性主基因遗传模型, 主基因遗传率为42.40%; 在QTL分析中, 用CIM和MIM共同检测到3个耐淹QTL, 分别位于A1、D1a和G连锁群上的Satt648~K418_2V、Satt531~A941V、Satt038~Satt275 (B53B~Satt038)区间, 表型贡献率为4.4%~7.6%。分离分析与QTL定位的结果相对一致, 可相互印证。     

大豆重组自交系群体荚粒性状的QTL分析

利用大豆重组自交系soy01群体中的255个家系进行2年田间试验，采用两种作图方法，寻找一粒荚、四粒荚、每荚粒数等5个荚粒性状稳定的QTL。结果表明，利用区间作图法，2年共找到24个荚粒性状QTL，解释的遗传变异为5%~80%；利用复合区间作图法，2年共找到27个荚粒性状QTL，解释的遗传变异为4%~73%。利用复合区间作图法，2年找到2个重复出现、稳定的四粒荚QTL和2个每荚粒数QTL，为大豆荚粒性状QTL的精细定位和分子标记辅助育种提供了基础和依据。     

Quantitative trait loci for early maturity and their potential in breeding for earliness in Brassica juncea

A doubled haploid population of Brassica juncea, developed from a cross between two parental lines differing for days to maturity, was used to study the efficiency of indirect selection for a primary trait through selection of secondary trait(s) over direct selection for the primary trait when quantitative trait loci information is available for both primary and secondary traits, and applied. Days to maturity was considered as primary trait, while days to first flowering, days to end of flowering, flowering period and plant height were considered as secondary traits. An RFLP linkage map was employed for QTL analysis of maturity and maturity-determinant traits, and a stable QTL B6 simultaneously affecting these two types of traits was identified. This linked QTL explained 11.7% phenotypic variation for days to maturity, 20.7% variation for days to first flowering, 24.3% variation for days to end of flowering and 14.4% variation for plant height. Phenotypic evaluation of maturity and/or maturity-determinant traits, viz. days to first flowering, days to end of flowering and plant height revealed that limited genetic advance for early maturity can be achieved through phenotypic selection of the primary and/or the secondary trait(s). However, the estimates of genetic advance for early maturity based on combined phenotypic evaluation and linked QTL data was found to be, at least, three times higher compared to genetic advance based on phenotypic evaluation only, demonstrating the potential of marker-assisted selection in breeding for early maturity in B. juncea.     

Genetic studies on saline and sodic tolerances in soybean

Salt-affected soils are generally classified into two main categories: saline and sodic (alkaline). Developing and using soybean (Glycine max (L.) Merr) cultivars with high salt tolerance is an effective way of maintaining sustainable production in areas where soybean growth is threatened by salt stress. Early classical genetics studies revealed that saline tolerance was conditioned by a single dominant gene. Recently, a series of studies consistently revealed a major quantitative trait locus (QTL) for saline tolerance located on linkage group N (chromosome 3) around the SSR markers Satt255 and Sat_091; other minor QTLs were also reported. In the case of sodic tolerance, most studies focused on iron deficiency caused by a high soil pH, and several QTLs associated with iron deficiency were identified. A wild soybean (Glycine soja Sieb. & Zucc.) accession with high sodic tolerance was recently identified, and a significant QTL for sodic tolerance was detected on linkage group D2 (chromosome 17). These studies demonstrated that saline and sodic tolerances were controlled by different genes in soybean. DNA markers closely associated with these QTLs can be used for marker-assisted selection to pyramid tolerance genes in soybean for both saline and sodic stresses.     

大豆不同环境下脂肪酸组分含量的QTL分析

大豆油的品质取决于脂肪酸各组分在大豆中的比例, 为发掘控制大豆5种脂肪酸含量的数量性状位点(QTL), 利用冀豆12和黑豆重组自交系群体构建遗传图谱, 采用Windows QTL Cartographer 2.5和QTL Network-2.0软件的CIM和MCIM法对大豆5种脂肪酸组分进行数量性状定位。结果表明,在石家庄和三亚各环境下共检测到16个QTL, 位于连锁群A2、B2、C2、F、G、I、L上。对2个环境联合分析, 检测到13个QTL, 其中9个用2种方法被检测到, 但这13个位点与环境互作的贡献率明显小于加性效应。其中在B2连锁群Satt168~Satt556控制硬脂酸的QTL Ste-1在河北石家庄和海南三亚均能被检测到, 贡献率均为12%, 在双尾群体和间隔挑选群体中也能检测到控制硬脂酸的QTL Ste-1, 说明这一QTL稳定存在于本组合群体中, 为今后大豆硬脂酸的QTL精细定位奠定了基础。     

大豆株高QTL发育动态分析

应用分子遗传连锁图谱和条件QTL定位方法对性状进行动态分析是发育遗传学新的研究方法。采用来自Charleston ×东农594的143个重组自交系（RILs），构建了一个20条连锁群的大豆分子遗传连锁图谱，以此为基础，采用复合区间作图法共定位了28个显著影响株高发育的非条件QTL，以条件分析方法和复合区间作图法相结合定位了21个影响株高发育的条件QTL。不同发育时期显著影响株高的QTL数目和遗传效应的变化，说明控制株高发育的数量基因位点是选择性表达的。因此，进行标记辅助选择时综合考虑不同发育时期表达的QTL，才能取得较好的效果。     

玉米开花期性状的QTL及杂种优势位点定位

开花期是玉米进化和适应过程中的重要性状,明确开花期杂种优势的遗传机制对培育适应不同生态区的优良玉米品种具有重要的意义。本研究利用以许178为受体,综3为供体构建的包含203个SSSL的单片段代换系群体及其与许178的测交群体,通过2年3个试点玉米开花期性状(散粉期、吐丝期和散粉至吐丝间隔)QTL和杂种优势位点(HL)分析,分别鉴定出40个开花期相关性状的QTL和37个开花期相关性状的HL。其中6个QTL和4个HL在3个地点被同时检测到。在所检测到的染色体区段中,11个区段同时包含调控开花期的QTL和HL。该研究为进一步解析玉米开花期遗传机制和开花期杂种优势的遗传机制提供了基础。     

QTL mapping of phosphorus deficiency tolerance in soybean (Glycine max L. Merr.)

Phosphorus (P) deficiency is a major abiotic stress that limits plant growth and crop productivity throughout the world. In the present study, 184 recombinant inbred line (RIL) families developed from soybean varieties Kefeng No. 1 and Nanong 1138-2 were used to identify quantitative trait loci (QTL) associated with P deficiency tolerance. Seven traits of plant height (HT), weight of fresh shoot (FSW), weight of fresh root (FRW), weight of dry root (DRW), length of main root (RL), phosphorus content in leaf (LP), phosphorus content in root (RP), were used as parameters to assess the phosphorus deficiency tolerance. The QTL mapping for the seven traits was performed using the program WinQTLCart. Seven QTLs were detected and mapped on two linkage groups for three traits of weight of fresh shoot, phosphorus contents in leaf and in root. The QTLs that had LOD scores more than three were detected for all of the three traits above. Most of the QTLs explained more than 10% of the total variation. The two QTLs for phosphorus content in leaf explained more than 20% of the total variation, respectively. Five QTLs were mapped on linkage group F2, and two on linkage F1. It was suggested that the genes related to phosphorus deficiency tolerance located on linkage group F in soybean.Contributed equally to this work.     

陆地棉早熟基因来源的遗传分析

棉花早熟性相关的性状多为复杂的数量性状,是由多个基因和环境共同作用的结果,其遗传基础研究比较困难。利用连锁作图和关联分析方法,对目标性状进行基因定位,找出与性状相关联的位点,为解析复杂性状的基因来源提供了新的手段。本文分别以早熟亲本新陆早8号和新陆早10号为母本,以陆地棉标准系TM-1为父本构建F₂作图群体,利用在亲本间筛选出的多态性SSR引物,通过JoinMap 3.0软件构建了2张遗传图谱,以Win QTLCart 2.5复合区间作图法在F₂~F_2:3中定位到控制全生育期、苗期、蕾期、花铃期和霜前铃数等早熟性相关性状的37个QTL。控制早熟性的有利等位基因多来自早熟祖先611波和金字棉。多数性状在两类材料中由不同的基因控制,且以加性遗传为主。在由43个陆地棉品种材料构成的自然群体中通过关联分析检测到54个与这些早熟性相关性状极显著关联的位点。研究表明连锁作图和关联分析的检测结果具有较高的可比性。本研究为早熟陆地棉聚合改良以及分子标记辅助育种打下了基础。