QTL and epistatic analyses of heterosis for seed yield and three yield component traits using molecular markers in rapeseed (Brassica napus L.)

Aiming to explore the basis of heterosis in rapeseed, QTLs for yield and three yield component traits were mapped and the digenic interactions were detected in an F₂ population derived from a cross between two elite rapeseed lines, SI-1300 and Eagle, in this study. Twenty-eight QTLs were detected for the four yield traits, with only two of them detected simultaneously in the Wuhan and Jingmen environments. Additive, partial dominance, dominance, and overdominance effects were all identified for the investigated traits. Dominance (including partial dominance) was shown by 55% of the QTLs, which suggests that dominance is a major genetic basis of heterosis in rapeseed. At the P ?? 0.01 level with 1000 random permutations, 108 and 104 significant digenic interactions were detected in Wuhan and Jingmen, respectively, for the four yield-related traits using all possible locus pairs of molecular markers. Digenic interactions, including additive by additive, additive by dominance, and dominance by dominance, were frequent and widespread in this population. In most cases (78.3%), the interactions occurred among marker loci for which significant effects were not detected by single-locus analysis. Some QTLs (57.1%) detected by single-locus analysis were involved in epistatic interactions. It was concluded that epistasis, along with dominance (including partial dominance), is responsible for the expression of heterosis in rapeseed.     

Identification of quantitative trait loci and epistatic interactions for plant height and heading date in rice

Appropriate heading date and plant height are prerequisites for attaining the desired yield level in rice breeding programs. In this study, we analyzed the genetic bases of heading date and plant height at both single- locus and two-locus levels, using a population of 240 F_2:3 families derived from a cross between two elite rice lines. Measurements for the traits were obtained over 2 years in replicated field trials. A linkage map was constructed with 151 polymorphic marker loci, based on which interval mapping was performed using Mapmaker/QTL. The analyses detected six QTLs for plant height and six QTLs for heading date; collectively the QTLs for heading date accounted for a much greater amount of phenotypic variation than did the QTLs for plant height. Two-way analyses of variance, with all possible two-locus combinations, detected large numbers (from 101 to 257) of significant digenic interactions in the 2 years for both traits involving markers distributed in the entire genome; 22 and 39 were simultaneously detected in both years for plant height and heading date, respectively. Each of the interactions individually accounted for only a very small portion of the phenotypic variation. The majority of the significant interactions involved marker loci that did not detect significant effects by single-locus analyses, and many of the QTLs detected by single-locus analyses were involved in epistatic interactions. The results clearly demonstrated the importance of epistatic interactions in the genetic bases of heading date and plant height. Received: 5 May 2001 / Accepted: 3 August 2001     

Partial Dominance,Overdominance and Epistasis as the Genetic Basis of Heterosis in Upland Cotton (Gossypium hirsutum L.)

Determination of genetic basis of heterosis may promote hybrid production in Upland cotton (Gossypium hirsutum L.). This study was designed to explore the genetic mechanism of heterosis for yield and yield components in F_{2: 3} and F_{2: 4} populations derived from a hybrid ‘Xinza No. 1’. Replicated yield field trials of the progenies were conducted in 2008 and 2009. Phenotypic data analyses indicated overdominance in F₁ for yield and yield components. Additive and dominance effects at single-locus level and digenic epistatic interactions at two-locus level were analyzed by 421 marker loci spanning 3814 cM of the genome. A total of 38 and 49 QTLs controlling yield and yield components were identified in F_{2: 3} and F_{2: 4} populations, respectively. Analyses of these QTLs indicated that the effects of partial dominance and overdominance contributed to heterosis in Upland cotton simultaneously. Most of the QTLs showed partial dominance whereas 13 QTLs showing overdominance in F_2:3 population, and 19 QTLs showed overdominance in F_2:4. Among them, 21 QTLs were common in both F_{2: 3} and F_{2: 4} populations. A large number of two-locus interactions for yield and yield components were detected in both generations. AA (additive × additive) epistasis accounted for majority portion of epistatic effects. Thirty three complementary two-locus homozygotes (11/22 and 22/11) were the best genotypes for AA interactions in terms of bolls per plant. Genotypes of double homozygotes, 11/22, 22/11 and 22/22, performed best for AD/DA interactions, while genotype of 11/12 performed best for DD interactions. These results indicated that (1) partial dominance and overdominance effects at single-locus level and (2) epistasis at two-locus level elucidated the genetic basis of heterosis in Upland cotton.     

Additive and over-dominant effects resulting from epistatic loci are the primary genetic basis of heterosis in rice

A set of 148 F9 recombinant inbred lines (RILs) was developed from the cross of an indica cultivar 93-11 and japonica cultivar DTT13,showing strong F1 heterosis.Subsequently,two backcross F1 (BCF1) populations were constructed by backcrossing these 148 RILs to two parents,93-11 and DT713.These three related populations (281BCF1 lines,148 RILs) were phenotyped for six yield-related traits in two locations.Significant inbreeding depression was detected in the population of RILS and a high level of heterosis was observed in the two BCF1 populations.A total of 42 main-effect quantitative trait loci (M-QTLs) and 109 epistatic effect QTL pairs (E-QTLs) were detected in the three related populations using the mixed model approach.By comparing the genetic effects of these QTLs detected in the RILs,BCF1 performance and mid-parental heterosis (HMp),we found that,in both BCF1 populations,the QTLs detected could be classified into two predominant types:additive and over-domlnant loci,which indicated that the additive and over-dominant effect were more important than complete or partially dominance for M-QTLs and E-QTLs.Further,we found that the E-QTLs detected collectively explained a larger portion of the total phenotypic variation than the M-QTLs in both RILs and BCF1 populations.All of these results suggest that additive and over-dominance resulting from epistatic loci might be the primary genetic basis of heterosis in rice.     

Analysis on additive effects and additive-by-additive epistatic effects of QTLs for yield traits in a recombinant inbred line population of rice

A linkage map consisting of 158 DNA markers were constructed by using a recombinant inbred line (RIL) population derived from the indica-indica rice cross Zhenshan 97B 2 Milyang 46. Quantitative trait loci (QTLs) conditioning grain yield and five yield component traits were determined at the one-locus and two-locus levels, and genotype-by-environment (GE) interactions were analyzed. Thirty-one QTLs were detected to have significant additive effects for yield traits, of which 12 also exhibited significant epistatic effects. Sixteen significant additive-by-additive (AA) interactions were detected, of which nine occurred between QTLs with own additive effects (M_epQTLs), four occurred between QTLs showing epistatic effects only (epQTLs), and three occurred between M_epQTLs and epQTLs. Significant GE interactions were found for six QTLs with additive effects and one AA interaction. Generally, the contributions to the phenotypic variation were higher due to QTL main effects than to epistatic effects. The detection of additive effects and AA effects of a QTL interfered with each other, indicating that the detection of QTLs with main effects, as well as the magnitude and directions of the additive effects, might vary depending on their interactions with other loci.     

水稻生物学产量及其构成性状的QTL定位

QTL的加性效应、加性×加性上位性效应及它们与环境的互作效应是数量性状的重要遗传分量.利用IR64/Azucena的125个DH品系为群体,分析了水稻生物学产量及其两个构成性状干草产量和谷粒产量的遗传组成.用基于混合模型的复合区间作图(MCIM)方法进行QTL定位.检测到12个位点有加性主效应,27个位点涉及双位点互作,18个位点存在环境互作.结果表明水稻生物学产量和它的两个构成性状普遍存在上位性效应和QE互作效应.此外,还探讨了性状间相关的遗传基础.发现4个QTLs和一对上位性QTLs可能与生物学产量与干草产量之间的正相关有关.3个QTL可能与干草产量与谷粒产量之间的负相关有关.这些结果可能部分地解释了这3个性状相关的遗传原因.通过对水稻生物学产量及其两个构成性状所定位QTL的分析,加深了对数量性状QTL的认识.首先,QTL的上位性效应和QE互作效应是普遍存在的;其次,QTL的多效性或紧密连锁可能是遗传相关的原因,当QTL对两个性状作用的方向相同时可导致正向遗传相关,反之则为负向遗传相关,当有些QTL表现为同向作用而另一些QTL表现为反向作用时,则可削弱性状间的遗传相关性;第三,复合性状的QTL效应可分解为其组成性状的QTL效应,如果QTL对各组成性状的效应方向相反而相互抵消,可使复合性状的QTL效应不易被检测;第四,加性效应的QTL常参预构成上位性效应,而具有上位性效应的QTL并非都有加性主效应,表明忽略上位性的QTL定位方法会降低检测QTL的功效;最后,鉴别不同类型的QTL效应有利于指导育种实践,选择主效QTL适用于多环境,QE互作QTL适用于特定环境,对上位性QTL应强调选择基因组合而并非单个基因.     

Identification of main effects, epistatic effects and their environmental interactions of QTLs for yield traits in rice

Quantitative trait loci (QTLs) controlling yield and yield components were identified by using a doubled haploid (DH) population of 120 lines from a sub-specific cross between ‘Samgang’ (Indica) and ‘Nagdong’ (Japonica). Main effects, epistatic effects, their environment interactions of QTLs were analyzed via mixed linear model approach across different environments. A total of 17 putative QTLs were identified on 8 chromosomes and five QTLs were detected over two years. 7 QTLs of main effects and 23 epistatic interactions were observed for five traits. Epistatic interactions played an important role in controlling the expression of yield related traits. The epistatic effects explained higher percentages of phenotype variation for panicles per plant, seed set percentage and yield. Significant QTL×environment (QE) interactions effects were identified for all traits, including 5 main effect QTLs. However, the present study failed to identify the significant interactions between epistatic loci containing main effect QTLs and the environment. The information provided in the present study could be used in the marker-assisted selection to enhance selection efficiency and to improve yield in rice.     

Characterization of the main effects,epistatic effects and their environmental interactions of QTLs on the genetic basis of yield traits in rice

Main effects, epistatic effects and their environmental interactions of QTLs are all important genetic components of quantitative traits. In this study, we analyzed the main effects, epistatic effects of the QTLs, and QTL by environment interactions (QEs) underlying four yield traits, using a population of 240 recombinant inbred lines from a cross between two rice varieties tested in replicated field trials. A genetic linkage map with 220 DNA marker loci was constructed. A mixed linear model approach was used to detect QTLs with main effects, QTLs involved in digenic interactions and QEs. In total, 29 QTLs of main effects, and 35 digenic interactions involving 58 loci were detected for the four traits. Thirteen QTLs with main effects showed QEs; no QE was detected for the QTLs involved in epistatic interactions. The amount of variations explained by the QTLs of main effect were larger than the QTLs involved in epistatic interactions, which in turn were larger than QEs for all four traits. This study illustrates the ability of the analysis to assess the genetic components underlying the quantitative traits, and demonstrates the relative importance of the various components as the genetic basis of yield traits in this population.     

Mapping QTL for Biomass Yield and Its Components in Rice (Oryza sativa L.)

Addicive effects, additive by additive epistatic effects, and their environmental interactions of QTLs are important genetic components of quantitative traits. Genetic architecture underlying rice biomass yield and its two component traits (straw yield and grain yield) were analyzed for a population of 125 DH lines from an inter-subspecific cross of IR64/Azucena. The mixed-model based composite interval mapping approach (MCIM) was used to detect QTLs, There were 12 QTLs detected with additive main effects, 27 QTLs involved in digenic interaction with aa and/or aae effects, and 18 QTLs affected by environments with ae and/or aae effects. It was revealed that epistatic effects and QE interaction effects existed on biomass yield and its component traits in rice. In addition, the genetic basis of relationships among these traits were investigated. Four QTLs and one pair of epistatic QTLs were detected to be responsible for the positive correlation between biomass yield and straw yield. Three QTLs might be responsible for the negative correlation between straw yield and grain yield. This result could partially explain the genetic basis of correlation among the three traits, and provide useful information for genetic improvement of these traits by marker-assisted selection.     

水稻加工品质数量性状基因座 (QTLs)分子定位研究

检测了Lemont/特青RI群体212个株系的糙米率（BR）,精米率（MR）和整精米率（HR）等3项加工品质性状,利用RFLP连锁图和线性模型的复合区间作图方法（QTLMapperV1.0)进行QTL定位研究。群体呈边境分布,双向超亲现象明显,HR较BR,MR变异范围更大并偏向低值方向;分别检测到1个MR,4个HR主效QTL,其中QHr6和QHr7等2个基因座具有较大遗传效应;分别检测到12对影响BR、5对影响MR,16对影响HR的上位性基因座,上位性效应的影响大于主效QTLs,不同性状或同一性状上位性效应通过共同的区间形成复杂的互相联系。     

Genetic variation underlying pod size and color traits of common bean depends on quantitative trait loci with epistatic effects

Common bean is an important vegetable legume in many regions of the world. Size and color of fresh pods are the key factors for deciding the commercial acceptance of bean as a fresh vegetable. The genetic basis of important horticultural traits of common bean is still poorly understood, which hinders DNA marker-assisted breeding in this crop. Here we report the identification of single-locus and epistatic quantitative trait loci (QTLs), as well as their environment interaction effects for six pod traits, namely width, thickness, length, size index, beak length and color, using an Andean intra-gene pool recombinant inbred line population from a cross between a cultivated common bean and an exotic nuña bean. The QTL analyses performed detected a total of 23 QTLs (single-locus QTLs and epistatic QTLs): five with only individual additive effects and six with only epistatic effects, while the remaining twelve showed both effects. These QTLs were distributed across linkage groups (LGs) 1, 2, 4, 6, 7, 8, 9, 10 and 11; particularly noteworthy are the QTLs for pod size co-located on LGs 1 and 4, indicative of tight linkage or genes with pleiotropic effects governing these traits. Overall, the results obtained showed that additive and epistatic effects are the major genetic basis of pod size and color traits. The mapping of QTLs including epistatic loci for the six pod traits evaluated provides support for implementing marker-assisted selection toward genetic improvement of common bean.     

Genetic Components of Heterosis for Seedling Traits in an Elite Rice Hybrid Analyzed Using an Immortalized F_2 Population

Utilization of heterosis has greatly contributed to rice productivity in China and many Asian countries. Superior hybrids usually show heterosis at two stages: canopy development at vegetative stage and panicle development at reproductive stage resulting in heterosis in yield. Although the genetic basis of heterosis in rice has been extensively investigated, all the previous studies focused on yield traits at maturity stage. In this study, we analyzed the genetic basis of heterosis at seedling stage making use of an "immortalized F2" population composed of 105 hybrids produced by intercrossing recombinant inbred lines(RILs) from a cross between Zhenshan 97 and Minghui 63,the parents of Shanyou 63, which is an elite hybrid widely grown in China. Eight seedling traits, seedling height, tiller number, leaf number, root number, maximum root length, root dry weight, shoot dry weight and total dry weight, were investigated using hydroponic culture. We analyzed single-locus and digenic genetic effects at the whole genome level using an ultrahigh-density SNP bin map obtained by population re-sequencing. The analysis revealed large numbers of heterotic effects for seedling traits including dominance, overdominance and digenic dominance(epistasis) in both positive and negative directions. Overdominance effects were prevalent for all the traits, and digenic dominance effects also accounted for a large portion of the genetic effects. The results suggested that cumulative small advantages of the single-locus effects and two-locus interactions, most of which could not be detected statistically, could explain the genetic basis of seedling heterosis of the F_1 hybrid.     

QTL analyses of heterosis for grain yield and yield-related traits in indica-japonica crosses of rice (Oryza sativa L.)

Two sets of rice materials, 166 RILs derived from a cross between Milyang 23 (Korean indica-type rice) and Tong 88-7 (japonica Rice), and BC₁F₁ hybrids derived from crosses between the RILs and the female parent, Milyang 23, were produced to identify QTLs for heterosis of yield and yield-related traits. The QTLs were detected from three different phenotype data sets including the RILs, BC1F1 hybrids, and mid-parental heterosis data set acquired from the definition of mid-parental heterosis. A total of 57 QTLs were identified for nine traits. Of eight QTLs detected for yield heterosis, five overlapped with other heterosis QTLs for yield-related traits such as spikelet number per panicle, days to heading, and spikelet fertility. Four QTLs for yield heterosis, gy1.1, py6, gy10, and py11, were newly identified in this study. We identified a total of 17 EpQTLs for yield heterosis that explain 21.4 ?? 59.0 % of total phenotypic variation, indicating that epistatic interactions may play an important role in heterosis.     

Dynamic analysis of QTLs on tiller number in rice (Oryza sativa L.) with single segment substitution lines

Twelve single segment substitution lines (SSSLs) in rice, which contain quantitative trait loci (QTLs) for tiller number detected previously, were used to study dynamic expression of the QTLs in this study. These SSSLs and their recipient, Hua-Jing-Xian 74 (HJX74), were used to produce 78 crossing combinations first, and then these combinations and their parents were grown in two planting seasons with three cropping densities. Tiller number was measured at seven developmental stages. QTL effects including main effects (additive, dominance and epistasis), QTL?×?season and QTL?×?density interaction effects were analyzed at each measured stage. The additive, dominant and epistatic effects of the 12 QTLs as well as their interaction effects with the seasons and with the densities all display dynamic changes with the development. Eight QTLs are detected with significant additive effects and/or additive?×?season and/or additive?×?density interaction effects at least at one developmental stage, and all QTLs have significant dominant and epistatic effects and/or interaction effects involved in. For most of the QTLs dominant effects are much bigger than additive effects, showing overdominance. Each QTL interacts at least with eight other QTLs. Additive and dominant effects of these QTLs are mostly positive while epistatic effects are negative and minor. Most of the QTLs show significant interactions with planting seasons and cropping densities, but the additive effects of QTLs Tn3-1 and Tn3-2, the dominant effects of QTL Tn7 and Tn8, and the epistatic effects of 14 pairs of QTLs are stable across seasons and the dominant effect of QTL Tn3-3 and the epistatic effects of QTL pairs Tn2-1/Tn6-2, Tn2-1/Tn9 and Tn3-3/Tn6-3 are nearly consistent across cropping densities. This paper is the first report of dynamics on dominances and epistasis of QTLs for tiller number in rice and provides abundant information, which is useful to improve rice tiller number via heterosis and/or QTL pyramiding.     

Genetic basis of heterosis for growth-related traits in Arabidopsis investigated by testcross progenies of near-isogenic lines reveals a significant role of epistasis

Epistasis seems to play a significant role in the manifestation of heterosis. However, the power of detecting epistatic interactions among quantitative trait loci (QTL) in segregating populations is low. We studied heterosis in Arabidopsis thaliana hybrid C24 x Col-0 by testing near-isogenic lines (NILs) and their triple testcross (TTC) progenies. Our objectives were to (i) provide the theoretical basis for estimating different types of genetic effects with this experimental design, (ii) determine the extent of heterosis for seven growth-related traits, (iii) map the underlying QTL, and (iv) determine their gene action. Two substitution libraries, each consisting of 28 NILs and covering approximately 61 and 39% of the Arabidopsis genome, were assayed by 110 single-nucleotide polymorphism (SNP) markers. With our novel generation means approach 38 QTL were detected, many of which confirmed heterotic QTL detected previously in the same cross with TTC progenies of recombinant inbred lines. Furthermore, many of the QTL were common for different traits and in common with the 58 QTL detected by a method that compares triplets consisting of a NIL, its recurrent parent, and their F(1) cross. While the latter approach revealed mostly (75%) overdominant QTL, the former approach allowed separation of dominance and epistasis by analyzing all materials simultaneously and yielded substantial positive additive x additive effects besides directional dominance. Positive epistatic effects reduced heterosis for growth-related traits in our materials.     

稻米粒形的QTL定位及上位性和QE互作分析

利用'广陆矮4号'×'佳辐占'水稻重组自交系构建了SSR标记的遗传图谱.联合2007年和2008年获得的两组稻米粒长(GL)、粒宽(GW)、长宽比(L/W)数据应用混合线性模型方法进行QTL定位,并作加性效应、加性×加性上位互作效应以及加性QTL、上位性QTL与环境的互作效应分析.结果显示;(1)在加性效应分析中两个群体共检测到4个控制粒长的QTL,4个控制粒宽的QTL,5个控制长宽比的QTL,贡献率分别为13.81%、15.36%和 16.29%.(2)在上位互作效应分析中两个群体共检测到2对控制粒长的互作QTL,1对控制粒宽的互作QTL,3对控制长宽比的互作QTL,贡献率分别为5.77%、2.59%和7.42%.(3)环境互作检测中,发现共有13个加性QTL和4对QTL的加性×加性上位性与环境产生了互作效应.结果表明,上位性效应和加性效应都影响稻米粒形遗传,QE互作效应也对粒形有着显著的影响.     

玉米株高和穗位高遗传基础的QTL剖析

利用玉米强优势组合(Mo17×黄早四)自交衍生的191个F2单株构建了由SSR和AFLP标记组成的分子连锁图谱.F2进一步自交产生的184个F2:3家系用于调查株高和穗位高的表型值.采用基于混合线性模型的复合区间作图法和相应的作图软件QTLmapper/V2.0,分别定位了7个株高和6个穗位高QTL;检测到18对控制株高和13对控制穗位高的上位性效应位点;同时发现了与环境存在显著互作的6个株高和8个穗位高单位点标记区域以及4对株高和4对穗位高上位性效应区域.分析了各种遗传因素在株高和穗位高遗传基础中的相对作用大小,指出了加性、显性和上位性是玉米株高和穗位高的重要遗传基础.并对所定位的QTL的真实性、株高和穗位高的关系以及研究结果对分子育种的启示予以讨论.     

家蚕茧质性状的QTL定位研究

采用QTLMapper 2．0 QTL作图软件,对F2群体的家蚕全茧量、茧层量、茧层率和蛹体重等性状进行了QTL定位分析,分别检测出7个、6个、2个、8个有显著效应分量的QTLs,分布于7个、5个、2个、7个不同的连锁群。控制全茧量、茧层量的QTLs一般存在复杂的上位性效应。对全茧量性状,有3对QTLs存在显著的加加上位性效应,其中1对还存在加显、显显互作;共有3个QTLs存在显著的显性效应,1个存在显著的加性效应。对茧层量QTLs,发现1对QTLs存在极显著的各项遗传效应,包括上位性效应;1对QTLs被检测到显著的显显互作,1个QTL具有显著的显性效应,并与另一个QTL存在显著的加加互作。茧层率、蛹体重主要受加性或显性的QTLs作用,没有发现茧层率QTLs的上位性效应,蛹体重的有效QTL大都呈现显著的负向显性效应,只有一对QTLs存在显著的加加上位性效应。第2、3、4、11、13、24、34、37、40连锁群是两个或多个性状QTLs分布的共同连锁群。全茧量和茧层量存在共同的QTL或染色体区域,育种上可通过适当选配,利用基因的互作效应,同步改良这两个性状。     

The genetic basis of stay-green in rice analyzed in a population of doubled haploid lines derived from an<Emphasis Type="Italic"> indica</Emphasis> by<Emphasis Type="Italic"> japonica</Emphasis> cross

Delayed leaf-senescence, or stay-green, has been regarded as a desired characteristic for the production of a number of crops including rice. In this study, we analyzed the genetic basis of stay-green using a population of 190 doubled haploid lines from the cross between an indica parent Zhenshan 97 and a stay-green japonica parent Wuyujing 2. The population was tested in replicated field trials in 2 consecutive years, and six traits were defined to evaluate the stay-green characteristics. A genetic linkage map with 179 SSR (simple sequence repeat) marker loci was constructed. The software QTLMapper, based on a mixed linear model approach, was applied to detect QTLs, epistatic effects and their environmental interactions for these traits. A total of 46 main-effect QTLs was detected for the six traits that can be localized to 25 chromosomal regions. The individual effects of all the QTLs were small. Fifty digenic interactions were resolved that involved 66 loci distributed on all 12 chromosomes. Environmental interactions were detected for 18 of the main-effect QTLs and 14 of the epistatic interactions. Collectively, the epistatic effects and QTL by year interactions accounted for large proportions of the phenotypic variations. The results also showed that most of the stay-green traits were negatively correlated with yield and its component traits. The implications of the results in crop improvement were discussed.Communicated by C. Möllers     

QTL mapping for yield and yield contributing traits in two mapping populations of bread wheat

In bread wheat, single-locus and two-locus QTL analyses were conducted for seven yield and yield contributing traits using two different mapping populations (P I and P II). Single-locus QTL analyses involved composite interval mapping (CIM) for individual traits and multiple-trait composite interval mapping (MCIM) for correlated yield traits to detect the pleiotropic QTLs. Two-locus analyses were conducted to detect main effect QTLs (M-QTLs), epistatic QTLs (E-QTLs) and QTL × environment interactions (QE and QQE). Only a solitary QTL for spikelets per spike was common between the above two populations. HomoeoQTLs were also detected, suggesting the presence of triplicate QTLs in bread wheat. Relatively fewer QTLs were detected in P I than in P II. This may be partly due to low density of marker loci on P I framework map (173) than in P II (521) and partly due to more divergent parents used for developing P II. Six QTLs were important which were pleiotropic/coincident involving more than one trait and were also consistent over environments. These QTLs could be utilized efficiently for marker assisted selection (MAS).