Dynamic analysis of QTL for plant height at different developmental stages in maize （Zea mays L.）

Plantheightisoneofimportantagronomictraitsinmaizebreeding.Inthepastfewyears,toincreasetheplantingdensityandpreventplantsfromlodging,studiesonthegeneticmechanismofplantheightweregivengreatattentionto.Sincethe1990s,molecularmarkershaveprovidedapowerfultooltostudythetraitofplantheightatthemolecularlevel[1—3].Butmostofresearchforplantheightonlyfocusedondataatmaturestage.Tillnow,about70genesorQTLshavebeenlocated[4].Moreover,somegeneshavebeenevencloned[5—7].Duringthevege-tativegrowthperiod,plant…     

Statistical method for mapping QTLs for complex traits based on two backcross populations

Most important agronomic and quality traits of crops are quantitative in nature.The genetic variations in such traits are usually controlled by sets of genes called quantitative trait loci (QTLs),and the interactions between QTLs and the environment.It is crucial to understand the genetic architecture of complex traits to design efficient strategies for plant breeding.In the present study,a new experimental design and the corresponding statistical method are presented for QTL mapping.The proposed mapping population is composed of double backcross populations derived from backcrossing both homozygous parents to DH (double haploid) or RI (recombinant inbreeding) lines separately.Such an immortal mapping population allows for across-environment replications,and can be used to estimate dominance effects,epistatic effects,and QTL-environment interactions,remedying the drawbacks of a single backcross population.In this method,the mixed linear model approach is used to estimate the positions of QTLs and their various effects including the QTL additive,dominance,and epistatic effects,and QTL-environment interaction effects (QE).Monte Carlo simulations were conducted to investigate the performance of the proposed method and to assess the accuracy and efficiency of its estimations.The results showed that the proposed method could estimate the positions and the genetic effects of QTLs with high efficiency.     

水稻DH群体剑叶宽度的QTL及其上位性和环境互作效应研究

利用基于混合模型的QTL定位方法,可以分析水稻DH群体在4个年份下剑叶宽度的QTL及其上位性效应和环境互作效应.结果发现,QTL除了具有自身的效应外,还可以参与上位性效应的形成,一个QTL可与多个QTL发生互作,可能起到诱发和修饰其他位点的作用.QTL与环境的互作效应以及上位性与环境的互作效应更多地被检测到,表明控制剑叶宽度的遗传因素易受环境影响.     

QTL mapping of the root traits and their correlation analysis with drought resistance using DH lines from paddy and upland rice cross

A Double Haploid (DH) population, 116 plants, derived from the cross between Japonica upland rice IRAT109 and paddy rice Yuefu, planted in PVC pipe under upland ecosystem in 2001 and 2002, was used in this study. Seven root traits, including basal root thickness (BRT), total root number (RN), maximum root length (MRL), root fresh weight (RFW), root dry weight (RDW), ratio of root fresh weight to shoot fresh weight (RFW/SFW) and ratio of root dry weight to shoot dry weight (RDW/SDW), were studied. Using index of drought resistance (IDR), the ratio of yield under upland ecosystem to yield under lowland ecosystem of DH lines, as the criteria of drought resistance, and correla-tion analysis between root traits and IDR, showed that BRT, MRL and RN were significantly correlated with IDR. High IDR lines had thicker BRT, longer MRL and less RN than low IDR lines. A molecular linkage map with 94 RFLP markers and 71 SSR markers covering 1535.1 cM was pro-duced. QTLs and G譋 interactions for BRT, RN, MRL, RFW, RDW, RFW/SFW and RDW/SDW were obtained based on the constructed molecular linkage map and soft-ware QTLmapper version 1.0. A total of 18 additive QTLs and 18 pairs of epistatic QTLs associated with root traits were detected. There were nine additive QTLs and two pairs of epistatic QTLs performed significant interactions with environment. Some QTLs with high general contribution and no G譋 interaction were obtained. Two pairs of epistatic QTLs mrl3 and mrl8, brt3 and brt11a controlling MRL and BRT had high general contributions of 21.51% and 13.03% respectively. An additive QTL and a pair of epistatic QTLs controlling RFW and RDW had high general contributions of 13.50% and 25.64% respectively. Marker assisted selec-tion (MAS) for rice drought resistance based on QTL with high general contribution, low G譋 interaction and tightly linkage with IDR were also discussed.     

玉米种皮和穗轴中总酚含量的QTL分析

利用黑玉米自交系SDM作为父本,与黄玉米自交系MO17杂交,构建F2∶3群体,对玉米种皮中总酚含量(PCP)和穗轴中的总酚含量(PCC)进行了QTL分析.结果表明:SDM种皮和穗轴中总酚含量均极显著高于MO17,种皮和穗轴中的总酚含量呈极显著正相关.与玉米种皮和穗轴中总酚含量相关的QTL各检测到4个,分布在第4,6,7,9,10共5条染色体上,增效基因均来自父本SDM.其中位于第6和第10染色体上的4个QTL对表型的贡献率最大.控制种皮中总酚含量的2个QTL PCP-6a和PCP-10b分别解释表型变异的13.00%和17.05%,所在的标记区间分别为mmc0523-umc2006和umc1506-bnlg1028,基因作用方式分别为部分显性和加性.控制穗轴中总酚含量的2个QTL PCC-6a和PCC-10a分别解释表型变异的12.20%和21.10%,所在的标记区间分别为umc1014-mmc0523和umc1506-bnlg1028,基因作用方式均为部分显性.位于第6和第10染色体上控制2个性状的QTL可能是一因多效的同一QTL,也可能是紧密连锁的不同QTL.     

Molecular dissection of plant height QTLs using recombinant inbred lines from hybrids between common wheat （Triticum aestivum L.） and spelt wheat （Triticum spelta L.）

In wheat, plant height is an important agronomic trait, and a number of quantitative trait loci (QTLs) controlling plant height have been located. In this study, using the conditional and unconditional QTL mapping methods, combined with data from five different growth stages over two years of field trials, the developmental behavior for plant height in wheat was dissected. Nine unconditional QTLs and 8 conditional QTLs were identified, of which 6 were detected by both methods. None of the 11 QTLs was detected at all of the 5 investigated developmental stages, but 7 QTLs were detected at certain stages in both years. Further analysis identified 9 unconditional QTLs at different stages, which could explain the phenotypic variation from 4.81% to 17.35%. It was noteworthy that one major QTL designated QHt-4B-2, which was located on chromosome 4B, was detected on May 18 and 25 in both years, and its genetic contributions to plant height ranged from 13.42% to 16.13%. Moreover, of the 8 conditional QTLs identified, six were detected in both years, in the order of QHt-3B→QHt-4B-1→QHt-4B-2→QHt-4D→QHt-5A and QHt-2B expressed at the same developmental stage. The results indicate that QTL expression during plant height development is selective and in a temporal order.     

QTL analysis of flag leaf in barley （Hordeum vulgate L.） for morphological traits and chlorophyll content

To understand genetic patterns of the morphological and physiological traits in flag leaf of barley, a double haploid （DH） population derived from the parents Yerong and Franklin was used to determine quantitative trait loci （QTL） controlling length, width, length/width, and chlorophyll content of flag leaves. A total of 9 QTLs showing significantly additive effect were detected in 8 intervals on 5 chromosomes. The variation of individual QTL ranged from 1.9% to 20.2%. For chlorophyll content expressed as SPAD value, 4 QTLs were identified on chromosomes 2H, 3H and 6H; for leaf length and width, 2 QTLs located on chromosomes 5H and 7H, and 2 QTLs located on chromosome 5H were detected; and for length/width, I QTL was detected on chromosome 7H. The identification of these QTLs associated with the properties of flag leaf is useful for barley improvement in breeding programs.     

Dissecting anxiety-related QTLs in mice by univariate and multivariate mapping

To detect genes underlying anxiety-related traits in mice,we performed univariate and multivariate QTL mapping analyses of phenotypes obtained from 71 mice of the BXD recombinant inbred (RI) strains (n=528 mice) and their parental strains (C57BL/6J and DBA/2J).Separate and joint mapping analyses were carried out using a linkage map composed of 506 simple sequence repeats (SSRs).The main QTL effects,interactions between pairs of QTLs (epistasis),and their environmental interactions were estimated.The results showed that anxiety-related traits were influenced by multiple QTLs (five main effect QTLs and three epistatic QTLs).Ten potential anxiety-related candidate genes within the QTL intervals on chromosomes 5,13 and 15 were identified.Some of these genes have been reported previously to be associated with the anxiety response.Based on our results,it is suggested that the multivariate QTL mapping approach improves the statistical power for detecting QTL and the precision of parameter estimation.Moreover,multivariate mapping can also detect pleiotropic QTL effects.     

玉米穗部性状的QTL定位

以玉米自交系L26和095组配的Fz世代为定位群体,采用SSR分子标记技术构建了包括98个位点的连锁图谱,结合F2穗部性状的鉴定结果,利用复合区间作图法对秃尖长等8个穗部性状进行基因定位,共检出21个QTL.其中穗长检测到3个QTL;穗粗、穗行数分别检测到2个QTL;行粒数检测到3个QTL;轴粗检测到2个QTL;200粒质量检测到3个QTL;穗粒质量检测到6个QTL;秃尖长没有检测到QTL.检出的21个QTL中,有10个QTL的解释变异率超过了20%,表现为主效QTL效应.研究还发现,穗部性状QTL在玉米10条染色体上分布不均匀,且成簇分布.该试验中检测到的21个QTL中,有10个影响不同性状的QTL位于3个染色体区域.各个QTL位点上起增、减效作用的等位基因在亲本间分布不均匀.     

水稻芒长及其分布特征相关QTL的定位

水稻芒的性状与驯化过程密切相关,具体表现为芒长短和芒的分布等．利用一个无芒品种越光（轮回亲本）和一个有芒品种Kasalath杂交产生的回交重组自交系群体,在上海、海南两地对芒长和芒的分布（芒出现比例）进行了相关分析及其相关基因（QTL）定位,结果发现这两个性状间存在显著的正相关．共检测到影响这两个性状的12个QTL,分布在6条染色体上的7个区域,贡献率介于2．90％-53．24％,其正效应大多来自有芒亲本Kasalath,但也有3个QTL正效应来自无芒亲本越光,并对这个现象产生的原因进行了讨论．本研究检测到的QTL及其两侧的分子标记可以用于水稻驯化的研究和理想型无芒品种分子辅助育种．     

Comparative mapping of rice root traits in seedlings grown in nutrient or non-nutrient solution

The component and amount of nutrient in the growth medium are the major factors affecting root growth.For the systematic dissection of root gene expression,evaluation of nutrient and non-nutrient solutions was conducted for their effect on root traits and quantitative trait loci(QTL)mapping.Three rice root parameters,maximum root length(MRL),root dry weight(RDW),and root/ shoot ratio of dry weight(RSR),were characterized within a double haploid(DH)population from a cross of ZYQ8(indica)and JX17(japonica).The value of the three root traits in two parents all decreased under the nutrient condition compared to those under the nonnutrient condition,of which RSR decreased up to 2.6-fold on average.In the DH population,more than 70 % lines in MRL,94 % lines in RDW,and all the lines in RSR were scored lower.In total,eight QTLs were identified in nutrient system(5 from JX17 alleles and 3 from ZYQ8 alleles)while five QTLs were detected in non-nutrient system(4 from JX17 alleles and 1 from ZYQ8 alleles).Of them,one QTL for RSR was shared by both culturing systems,seven QTLs were specific in nutrient system and the other four QTLs were specific in non-nutrient system.All 13 QTLs were distributed over 7 rice chromosomes-2,3,4,5,6,9 and 10,respectively.     

Identification and mapping of genes for improving yield from Chinese common wild rice (O. rufipogon Griff.) using advanced backcross QTL analysis

To identify useful genes from wild rice which have been lost or weakened in cultivated rice has become more and more important for modern breeding strategy. In this study, a BC4 population derived from 94W1, an acces-sion of common wild rice (Oryza rufipogon Griff.) from Dongxiang in Jiangxi Province of China, as the donor, and a high-yielding Indica cultivar (O. sativa L.), "Guichao 2", as the recipient, was used to identify quantitative trait loci (QTL) associated with yield and its components. Based on the analysis for the genotype of BC4F1 population with 87 SSR markers distributed throughout the genome and investigation of the plant height, yield and yield components of BC4F2, a total of 52 QTLs, were detected. Of 7 QTLs associated with grain yield per plant, 2 QTLs on chro-mosome 2 and chromosome 11 for grain yield, explaining 16% and 11% of the phenotypic variance respectively, were identified. The alleles from Dongxiang common wild rice in those two loci could increase the yield of "Guichao 2" by 25.9% and 23.2% respectively. The QTL on chromosome 2 increasing grain yield of cultivar is actually a major gene, which did not coincide with any previously published QTLs in rice.     

Quantitative trait loci （QTLs） for quality traits related to protein and starch in wheat

Quality traits in wheat （Triticum aestirum L.） were studied by quantitative trait locus （QTL） analysis in a recombinant inbred line （RIL） population, a set of 131 lines derived from Chuan 35050 × Shannong 483 cross （ChSh）. Grains from RILs were assayed for 21 quality traits related to protein and starch. A total of 35 putative QTLs for 19 traits with a single QTL explaining 7.99-40.52% of phenotypic variations were detected on 10 chromosomes, 1D, 2A, 2D, 3B, 3D, 5A, 6A, 6B, 6D, and 7B. The additive effects of 30 QTLs were positive, contributed by Chuan 35050, the remaining 5 QTLs were negative with the additive effect contributed by Shannong 483. For protein traits, 15 QTLs were obtained and most of them were located on chromosomes 1 D, 3B and 6D, while 20 QTLs for starch traits were detected and most of them were located on chromosomes 3D, 6B and 7B. Only 7 QTLs for protein and starch traits were co-located in three regions on chromosomes 1D, 2A and 2D. These protein and starch trait QTLs showed a distinct distribution pattern in certain regions and chromosomes. Twenty-two QTLs were clustered in 6 regions of 5 chromosomes. Two QTL clusters for protein traits were located on chromosomes 1D and 3B, respectively, three clusters for starch traits on chromosomes 3D, 6B and 7B, and one cluster including protein and starch traits on chromosome 1D.     

Resolution of quantitative traits into Mendelian factors by using a complete linkage map of restriction fragment length polymorphisms

The conflict between the Mendelian theory of particulate inheritance and the observation of continuous variation for most traits in nature was resolved in the early 1900s by the concept that quantitative traits can result from segregation of multiple genes, modified by environmental effects. Although pioneering experiments showed that linkage could occasionally be detected to such quantitative trait loci (QTLs), accurate and systematic mapping of QTLs has not been possible because the inheritance of an entire genome could not be studied with genetic markers. The use of restriction fragment length polymorphisms (RFLPs) has made such investigations possible, at least in principle. Here, we report the first use of a complete RFLP linkage map to resolve quantitative traits into discrete Mendelian factors, in an interspecific back-cross of tomato. Applying new analytical methods, we mapped at least six QTLs controlling fruit mass, four QTLs for the concentration of soluble solids and five QTLs for fruit pH. This approach is broadly applicable to the genetic dissection of quantitative inheritance of physiological, morphological and behavioural traits in any higher plant or animal.     

Correlation analysis and QTL identification for canopy temperature, leaf water potential and spikelet fertility in rice under contrasting moisture regimes

Canopy temperature (CT), leaf water potential(LWP) and spikelet fertility (SF) of a set of RILs (F9) from the cross between Zhenshan97B and IRAT109 were investigated under two soil moisture regimes in a drought screen facility. In water stress condition, CT was negatively correlated with SF (r = -0.2867) and LWP (r = -0.2740), and LWP was positively associated with SF (r = 0.1696). These results indicated that the plant with high drought tolerant ability could maintain higher LWP and lower CT, leading to highe rSF. A total of 44 main effect QTLs were associated with CT,LWP and SF. The accumulated contributions of QTLs for CT,LWP and SF were 87.85%, 15.06%, 79.46% under well water condition and 72.61%, 87.68%, 33.29% under stress condition, respectively. Totally 45 pairs of digenic interactions were detected. The accumulated contribution of digenic epistasis on CT, LWP and SF were 55.69%, 47.15%, 48.15% under well water condition and 53.44%, 57.94%, 54.62% under water stress condition. Compared with other drought tolerance QTL mapping researches in rice, 19 main effect QTLs were found to be located at the same or closely conjoint regions.     

影响小麦加工品质数量性状位点的研究

以M5和M16为亲本构建的重组自交系为材料,对影响小麦加工品质性状的数量性状位点（QTL）进行了研究．检测了群体中籽粒硬度、蛋白质含量、SDS沉降值、拉伸面积等品质性状,它们在群体中呈近似正态的连续分布,为多基因控制的数量性状．利用43个SSR标记和42个AFLP标记构建了相关染色体的分子连锁图．在1A、5D、6D染色体上分别检测到与籽粒硬度相关的QTL各1个,其中位于染色体5D上的Xgwn190对表型变异的贡献率最大,达62．5％,为主效基因;在1B和6A染色体上分别检测到与蛋白质含量相关的QTL各1个,它们对表型变异的贡献率分别为13．2％和15．6％;在染色体1B和3B上分别检测到与SDS沉降值相关的QTL各1个,贡献率最大（10．2％）的一个QTL位于3B染色体上靠近E37M61—286的区域;在1A、3B、5D染色体上分别检测到与拉伸面积相关的QTL各1个,其中5D染色体上的Xgwn190对表型的贡献率最大,为11．5％．     

水稻苗期形态性状的QTL定位

采用苗期形态性状上有着明显差异的粳稻Asominori和籼稻IR24杂交产生的重组自交系（RILs）为材料,经营养液培养后,对水稻苗期形态相关的6个数量性状：最大根长（MRL）、苗高（SH）、根干重（RDW）、茎（叶）干重（SDW）、总重（TW）以及根/茎（叶）干重比（RSR）进行了数量性状位点（QTL）定位分析.结果共检测到影响6个形态性状的8个QTL位点,分别位于水稻的第2,4,10和11染色体上,其贡献率为9.79%～22.90%.并发现位于第4染色体上的控制根/茎（叶）干重比的qRSR-4和根干重的qRDW-4的2个位点以及位于第10染色体上的控制茎（叶）干重的qSDW-10和总重的qTW-10的2个位点分别位于同一分子标记区间.研究结果对水稻苗期形态性状QTL的精细定位及培育理想株型具有一定的应用价值.     

Molecular dissection of genetic basis of significant correlation among five morphological traits in rice (Oryza sativa L.)

To enhance understanding of the genetic basis of trait correlation in rice, a recombinant inbred line (RIL) population (F₆ and F₇) from a cross between Zhenshan97 and HR5 was employed to identify main quantitative trait loci (QTLs) and epistatic QTL (E-QTL). Highly significant positive correlations were detected among five traits of heading date (HD), plant height (PH), panicle length (PL), flag leaf length (FLL) and flag leaf width (FLW) in 2 environments. Four to 8 main QTLs were detected for an individual trait. No E-QTL was detected for PH. One, 4, 4 and 5 E-QTLs were detected for FLL, HD, FLW and PL, respectively. Each E-QTL individually explained less than 3% of trait variation except E-QFll1. Comparison of QTL results was made in order to dissect the genetic basis of trait correlation. We found that main QTLs with pleiotropic effects and QTL clusters were the main genetic basis of trait correlation. No E-QTL had pleiotropic effects. E-QTL played an important role in the genetic basis of individual trait, but it made a little contribution to trait correlation.     

Genetic analysis of rice allelopathy

A double haploid population derived from anther culture of ZYQ8/JX17,a typical indica and japonica hybrid,was used in this study.The inhibited effect of water-soluble extract of 123 DH pure lines leaves on the lettuce roots growth was investigated,and the QTLs analysis of rice allelopathy was carried out.Totally,four QTLs related to rice allelopathy were detected,and they were on chromosomes,3,9,10 and 12,respectively.The LOD scores were 3.40,2.68,2.75 and 3.08,respectivel.Among them,additive effects of the QTLs on chromosomes 3 and 10 were 1.65 and 1.43,on chromosomes 9 and 12 were-1.44 and -1.58 respectively.Allelopathy characteristics of another three common rice varieties were also studied.