不同发育阶段大豆株高和茎粗QTL的动态分析

利用中豆29×中豆32的重组自交系,以复合区间作图法对不同发育阶段的大豆株高和茎粗同时进行非条件和条件QTL定位,在11个连锁群检测到18个株高QTL,在9个连锁群检测到19个茎粗QTL。不同发育时期影响大豆株高和茎粗QTL的数量、加性效应和贡献率均不相同,QTL表达具有时序性和选择性,有些QTL仅表达1次,有些可多次连续表达。有3个株高QTL和1个茎粗QTL在3个年度重复表达,有6个株高QTL和2个茎粗QTL在2个年度重复表达。F连锁群上株高和茎粗QTL存在共位性,R1～R4期均有株高和茎粗QTL同时表达,但株高和茎粗QTL的增效基因不同,株高QTL表达次数多而茎粗QTL表达次数较少,前期(V4～R3)QTL表达数量多而后期(R4～R5)表达数量较少。株高和茎粗QTL的动态变化与表型相关分析结果一致,对于适期选择粗秆抗倒的高产材料具有指导作用。     

Identification of genomic regions determining flower and pod numbers development in soybean （Glycine max L.）

Flower and pod numbers per plant are important agronomic traits underlying soybean yield.So far quantitative trait loci (QTL) detected for flower and pod-related traits have mainly focused on the final stage,and might therefore have ignored genetic effects expressed during a specific developmental stage.Here,dynamic expressions of QTL for flower and pod numbers were identified using 152 recombinant inbred lines (RILs) and a linkage map of 306 markers.Wide genetic variation was found among RILs;17 unconditional and 18 conditional QTL were detected for the two traits at different developmental stages over two years.Some QTL were detected only at one stage and others across two or more stages,indicating that soybean flower and pod numbers development may be governed by time-dependent gene expression.Three main QTL (qfn-Chr18-2,qfn-Chr20-1,and qfn-Chr19) were detected for flower number,and two main QTL (qpn-Chr11 and qpn-Chr20) were detected for pod number.The phenotypic variation explained by them ranged from 6.1% to 34.7%.The markers linked to these QTL could be used in marker-assisted selection for increasing soybean flower and pod numbers,with the ultimate aim of increasing soybean yield.Comparison of the QTL regions for flower and pod numbers traits with the related genes reported previously showed that seven and four related genes were located in the QTL regions of qfn-Chr11 and qfn-Chr19,respectively.Tbese results provide a basis for fine mapping and cloning of flower and pod development-related genes.     

Dynamic QTL analysis of the Na+ content,K+ content,and Na+/K+ ratio in rice roots during the field growth under salt stress

Rice (Oryza sativa L.) is seriously impacted by global soil salinization. To determine the quantitative trait loci (QTLs) related to salt tolerance in rice roots, F_2:3 and BC₁F_2:3 populations derived from a cross between the cv. Dongnong 425 of high quality and yield and the salt-tolerant cv. Changbai 10, were studied at different development stages. Two genetic linkage maps of F_2:3 and BC1F_2:3 populations were constructed. A 66 mM NaCl solution was used to irrigate the field and to analyze the dynamic QTL of some rice root traits. Using unconditional and conditional QTL mapping methods, 30 unconditional QTLs and 16 conditional QTLs related to the 6 root traits were detected on the 9 rice chromosomes during different developmental stages. Fourteen pairs of unconditional and conditional QTLs were detected at the identical developmental stage in the identical population. A number of QTLs were detected at different developmental stages, however, many did not appear at the last stage. Remarkably, qRKC1 appeared continuously at multiple stages in both the populations suggesting its key role in regulating the salt tolerance of rice roots.     

Quantitative trait loci underlying the development of seed composition in soybean (Glycine max L. Merr.).

One hundred and forty-three F2:7 recombinant inbred lines (RILs) developed from the cross of soybean cultivars 'Charleston' and 'Dongnong 594' were analyzed for the quantitative trait loci (QTLs) underlying protein or oil content at 6 different developmental stages by composite interval mapping with a mixed genetic model. The genotype x environment (GxE) interactions of the QTLs were also evaluated. Nineteen (2004) and 33 (2005) unconditional QTLs underlying seed protein or oil content at the different developmental stages were mapped onto 8 and 9 linkage groups, respectively. The proportion of phenotypic variation explained by these QTLs ranged from 6.26% to 30.52% and from 5.38% to 28.47%, respectively. Fourteen (2004) and 21 (2005) conditional QTLs underlying seed protein or oil content were mapped onto 5 and 8 linkage groups, respectively. The proportion of phenotypic variation explained by these QTLs ranged from 2.97% to 29.68% and from 5.42% to 31.96%, respectively. The numbers and types of QTLs and the genetic effect for the two traits were different at each developmental stage. However, several genomic regions that simultaneously control the development of both traits were detected. The genetic effect on protein content and oil content was opposite for loci in the marker interval Satt335-SSatt334, reflecting a negative correlation of protein content and oil content. A G x E interaction effect of some QTLs underlying protein or oil content at different growth periods was observed.     

Dynamic QTL Analysis of Rice Protein Content and Protein Index Using Recombinant Inbred Lines

Protein content (PC) and protein index (PI) play important roles in determining nutritional quality in rice (Oryza sativa L.). We used 71 lines derived from “Asominori/IR24” to analyze the developmental behavior of PC and PI through unconditional and conditional QTL mapping methods. In all, 10 unconditional QTLs and 6 conditional QTLs for PC, and 11 unconditional QTLs and 9 conditional QTLs for PI, were identified at four stages of grain filling. More were identified in the first three stages than at the final stage. Temporal patterns of gene expression for PC and PI differed over time, with several QTLs being expressed across two or three stages but many being expressed at only one stage. Some of these QTLs were closely linked with maturity QTLs reported previously. Many QTLs for PC and PI were co-localized, supporting the significant correlation found between PC and PI. Our results suggest that dynamic QTL mapping might be a valid means for revealing more genetic information about protein accumulations during seed development.     

Quantitative trait loci analysis for the developmental behavior of tiller number in rice (Oryza sativa L.)

 A doubled-haploid rice population of 123 lines from Azucena/IR64 was used for analyzing the developmental behavior of tiller number by conditional and unconditional QTL mapping methods. It was indicated that the number of QTLs significantly affecting tiller number was different at different measuring stages. Many QTLs controlling tiller growth identified at the early stages were undetectable at the final stage. Only one QTL could be detected across the whole growth period. By conditional QTL mapping, more QTLs for tiller number could be detected than that by unconditional mapping. The temporal patterns of gene expression for tiller number could be different at different stages. Even an individual gene or genes at the same genomic region might have opposite genetic effects at various growth stages. Received: 7 July 1997 / Accepted: 10 February 1998     

红麻株高与茎粗性状的动态发育遗传分析

红麻株高、茎粗是两个最重要的动态发育的经济性状.发育遗传学已探明生物个体的不同发育阶段基因是按一定的时空秩序有选择地表达的.采用数量性状的加性显性遗传模型和发育遗传模型,分析了红麻株高与茎粗在不同生长发育时期的遗传规律,即估算了株高与茎粗在某一生长发育时间(0→t)或某一特定生长发育时间段(t-1→t)的遗传效应.结果表明,株高在不同生长发育时期的非条件、条件遗传效应两者基本一致,均以显性效应为主,加性效应较弱,但条件遗传效应的分析与实际更为相符;茎粗的非条件、条件遗传效应表现一致,各个生长发育时期均未检测到加性效应,而显性效应均达显著、极显著水平;在各个不同生长发育时期中,株高与茎粗均在7月28日～8月9日(旺长期)、9月2～14日(纤维累积期)之间基因表达较活跃.株高与茎粗相关分析表明,两者在各个生长发育阶段均不存在加性相关,但8月21日之后的各期株高与茎粗之间均为显著或极著显正相关,该时期后进行相关选择比较有效.     

Dynamic QTL analysis of linolenic acid content in different developmental stages of soybean seed

Linolenic acid (LN) in soybean (Glycine max L. Merr.) seed mainly contributes to the undesirable odors and flavors commonly associated with poor oil quality. LN deposition at various stages of soybean seed development had not been reported by 2010. The objects of this study were (1) to identify and measure quantitative trait loci (QTL) underlying LN content and (2) to estimate the QTL effects expressed from earlier seed developmental stages to drying seed of soybean. One hundred and twenty-five F_5:8 and F_5:9 recombinant inbred lines derived from the cross of soybean cultivars ‘Hefeng 25’ and ‘Dongnong L5’ were used for the identification of QTL underlying LN content from the 37 day (D) to 86D stages after flowering, at Harbin in 2008 and 2009. QTL × Environment interactions (QE) effects were evaluated using a mixed genetic model (Zhu in J Zhejiang Univ (Natural Science) 33:327–335, 1999). Twelve unconditional QTL and 12 conditional QTL associated with LN content were identified at different developmental stages. Most of the QTL explained <10% of phenotypic variation of LN content. Unconditional QTL QLNF-1, QLNC2-1, QLND1b-1, QLNA2-1 and QLNH-1 influenced LN content across different development stages and environments. Conditional QTL QLNF-1, QLNC2-1 and QLNH-1 were identified in multiple developmental stages and environments. Conditional and unconditional QTL clustered in neighboring intervals on linkage groups A2, C2 and D1b. Ten QTL with conditional additive main effects (a) and/or conditional additive × environment interaction effects (ae) at specific developmental stage were identified on nine linkage groups. Of them, six QTL only possessed additive main effects and seven QTL had significant ae effects in different developmental stages. A total of 13 epistatic pairwise QTL were identified by conditional mapping in different developmental stages. Two pairs of QTL only showed aa effects and five pairs of QTL only showed aae effects at different developmental stages. QTL with aa effects, as well as their environmental interaction effects, appeared to vary at different developmental stages.     

Unconditional and conditional QTL mapping for the developmental behavior of tiller number in rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

A single segment substitution population of 26 lines and their recipient parent Hua-jing-xian 74 (HJX74) were selected as experimental materials for analyzing the developmental behavior of tiller number in rice. By the unconditional QTL (quantitative trait locus) mapping method, a total number of 14 SSSLs were detected with QTLs controlling rice tiller number. The number of QTLs significantly affecting tiller number and their effect values estimated differed across measuring stages. More QTLs could be detected based on time-dependent measures of different stages. By the conditional QTL mapping method, it is possible to reveal net expression of gene in a time interval. 14 QTLs on tiller number expressed their effects in dynamic patterns of themselves during whole ontogeny. They exhibited mainly negative effects within 7 days after transplanting. During 7–21 days, QTLs were in active status and expressed larger positive effects. In the mid-period of 21–35 days, they had opposite genetic effects to wither tillers. Since then these QTLs expressed positive effects again to cause the appearance of noneffective tillers. The dynamics of QTL effects was in agreement with the actual change of tillers. Mapping QTL combining unconditional with conditional analysis for time-dependent measures is helpful to understand roundly the genetic bases for the development of quantitative traits.     

Identification of QTLs for seed and pod traits in soybean and analysis for additive effects and epistatic effects of QTLs among multiple environments

Soybean seed and pod traits are important yield components. Selection for high yield style in seed and pod along with agronomic traits is a goal of many soybean breeders. The intention of this study was to identify quantitative trait loci (QTL) underlying seed and pod traits in soybean among eleven environments in China. 147 recombinant inbred lines were advanced through single-seed-descent method. The population was derived from a cross between Charleston (an American high yield soybean cultivar) and DongNong594 (a Chinese high yield soybean cultivar). A total of 157 polymorphic simple sequence repeat markers were used to construct a genetic linkage map. The phenotypic data of seed and pod traits [number of one-seed pod, number of two-seed pod, number of three-seed pod, number of four-seed pod, number of (two plus three)-seed pod, number of (three plus four)-seed pod, seed weight per plant, number of pod per plant] were recorded in eleven environments. In the analysis of single environment, fourteen main effect QTLs were identified. In the conjoint analysis of multiple environments, twenty-four additive QTLs were identified, and additive QTLs by environments interactions (AE) were evaluated and analyzed at the same time among eleven environments; twenty-three pairs of epistatic QTLs were identified, and epistasis (additive by additive) by environments interactions (AAE) were also analyzed and evaluated among eleven environments. Comparing the results of identification between single environment mapping and multiple environments conjoint mapping, three main effect QTLs with positive additive values and another three main effect QTLs with negative additive values, had no interactions with all environments, supported that these QTLs could be used in molecular assistant breeding in the future. These different effect QTLs could supply a good foundation to the gene clone and molecular asisstant breeding of soybean seed and pod traits.     

大豆小区产量及其相关性状QTL间的上位性和环境互作效应

以晋豆23栽培大豆(Glycine max)为母本、灰布支黑豆(ZDD2315,半野生大豆)为父本衍生出447个RIL群体,通过构建SSR遗传图谱及利用混合线性模型分析方法,对2年大豆小区产量及主要植物学性状进行QTL定位,并作加性效应、加性×加性上位互作效应及环境互作效应分析。结果显示,共检测到12个与小区产量、单株粒重、单株茎重、单株粒茎比、有效分枝、主茎节数、株高和结荚高度相关的QTL,分别位于A1、A2、H_1、I、J_2和M连锁群上。其中小区产量、株高、单株粒重、有效分枝和主茎节数均表现为遗传正效应,即增加其性状的等位基因来源于母本晋豆23。同时,检测到11对影响小区产量、单株粒重、单株茎重、株高和结荚高度的加性×加性上位互作效应及环境互作效应的QTL,发现22个QTL与环境存在互作。实验结果表明,上位效应和QE互作效应对大豆小区产量及主要农艺性状的遗传影响很大。进行大豆分子标记辅助育种时,既要考虑效应起主要作用的QTL,又要注重上位性QTL,这样有利于性状的稳定表达和遗传。     

Quantitative trait loci associated with oligosaccharide and sucrose contents in soybean (Glycine max L.)

Oligosaccharides and sucrose are very important nutritional components in soybean seeds. However, little information is available about their inheritance. We used molecular markers to identify the genomic regions significantly associated with the quantitative trait locus (QTL) that controls oligosaccharide and sucrose contents in segregating F_2:10 Rl lines. Two related, but independent, QTLs were identified for oligosaccharides — near marker satt546 on linkage group (LG) D1b+W and satt278 on LG L. Four others, for sucrose content, were located at LG B1 (satt197), D1b+W (satt546), and L (satt523 and satt278). Finally, we found two common QTLs, on LG D1b+W and L, that are associated with both oligosaccharides and sucrose.     

QTL mapping of ten agronomic traits on the soybean (<Emphasis Type="Italic">Glycine max</Emphasis> L. Merr.) genetic map and their association with EST markers

A set of 184 recombinant inbred lines (RILs) derived from soybean vars. Kefeng No.1 × Nannong 1138-2 was used to construct a genetic linkage map. The two parents exhibit contrasting characteristics for most of the traits that were mapped. Using restricted fragment length polymorphisms (RFLPs), simple sequence repeats (SSRs) and expressed sequence tags (ESTs), we mapped 452 markers onto 21 linkage groups and covered 3,595.9 cM of the soybean genome. All of the linkage groups except linkage group F were consistent with those of the consensus map of Cregan et al. (1999). Linkage group F was divided into two linkage groups, F1 and F2. The map consisted of 189 RFLPs, 219 SSRs, 40 ESTs, three R gene loci and one phenotype marker. Ten agronomic traits—days to flowering, days to maturity, plant height, number of nodes on main stem, lodging, number of pods per node, protein content, oil content, 100-seed weight, and plot yield—were studied. Using winqtlcart, we detected 63 quantitative trait loci (QTLs) that had LOD>3 for nine of the agronomic traits (only exception being seed oil content) and mapped these on 12 linkage groups. Most of the QTLs were clustered, especially on groups B1 and C2. Some QTLs were mapped to the same loci. This pleiotropism was common for most of the QTLs, and one QTL could influence at most five traits. Seven EST markers were found to be linked closely with or located at the same loci as the QTLs. EST marker GmKF059a, encoding a repressor protein and mapped on group C2, accounted for about 20% of the total variation of days to flowering, plant height, lodging and nodes on the main stem, respectively.Communicated by H.F. LinskensW.-K. Zhang, Y.-J. Wang and G.-Z. Luo contributed equally to this investigation.     

Molecular dissection of developmental behavior of tiller number and plant height and their relationship in rice (Oryza sativa L.)

Plant height and tiller number are two important characters related to yield in rice (Oriza sativa L.). Zhenshan97 x Minghui63 recombinant inbred lines were employed to dissect the genetic basis of development of plant height and tiller number using conditional and unconditional composite interval mapping approaches. The traits were normally distributed with transgressive segregation in both directions. Increasingly negative correlations were observed between tiller number and plant height at five consecutive growth stages. A total of 23 and 24 QTL were identified for tiller number and plant height, respectively. More QTL were detected by conditional mapping than by conventional mapping. Different QTL/genes apparently controlled the traits at different developmental stages. Three genomic regions were identified as putative co-located QTL, which showed opposite additive effects on tiller number and plant height. Furthermore, in the period reaching maximum tiller number, the expression of QTL for tiller number was active, whereas that of QTL for plant height was inactive. These facts provided a possible genetic explanation for the negative correlations between the traits. The research demonstrates conditional mapping to be superior to conventional mapping for this type of research. Implications of the results for hybrid rice improvement are discussed.     

Quantitative trait locus analysis of multiple agronomic traits in the model legume Lotus japonicus.

The first quantitative trait locus (QTL) analysis of multiple agronomic traits in the model legume Lotus japonicus was performed with a population of recombinant inbred lines derived from Miyakojima MG-20 x Gifu B-129. Thirteen agronomic traits were evaluated in 2004 and 2005: traits of vegetative parts (plant height, stem thickness, leaf length, leaf width, plant regrowth, plant shape, and stem color), flowering traits (flowering time and degree), and pod and seed traits (pod length, pod width, seeds per pod, and seed mass). A total of 40 QTLs were detected that explained 5%-69% of total variation. The QTL that explained the most variation was that for stem color, which was detected in the same region of chromosome 2 in both years. Some QTLs were colocated, especially those for pod and seed traits. Seed mass QTLs were located at 5 locations that mapped to the corresponding genomic positions of equivalent QTLs in soybean, pea, chickpea, and mung bean. This study provides fundamental information for breeding of agronomically important legume crops.     

QTLs conditioning early growth in a soybean population segregating for growth habit

 There are both economic and environmental reasons for reducing the use of herbicides for weed control in soybean [Glycine max (L.) Merr.] fields. Optimizing crop competitiveness can reduce reliance on chemical weed control. Fast and vigorous early growth and rapid canopy development can be effective in suppressing weed infestation of crop plants. The purposes of this study were to identify and molecularly map the quantitative trait loci (QTLs) conditioning soybean plant height and canopy width during the early vegetative stages of soybean growth. A restriction fragment length polymorphism (RFLP) linkage map was created using 142 markers and 116 F₂-derived lines from a cross of ‘S100’בTokyo’. The parents and the 116 F₂-derived lines were evaluated in the greenhouse and in the field at Athens, Ga., in 1996 and 1997. Combined over environments, Tokyo averaged 41 and 17% taller plants than S100 at the V7 and V10 stages of development. Transgressive segregation was observed among the progeny at both stages. Based on single-factor analysis of variance (ANOVA), three and four independent RFLP loci were associated with plant height at the V7 and V10 stages, respectively. All three loci detected [on linkage groups (LGs) C2 and F, and unlinked] at the V7 stage were also detected at the V10 stage along with one additional independent locus on LG E. The Tokyo allele contributed to increased plant height at all loci except at the unlinked locus. Three QTLs (on LGs C2, E, and F) were consistent across environments, three (on LGs C2 and F, and unlinked) were consistent across stages of plant development, and two (on LGs C2 and F) were consistent both across environments and stages of plant development. Within each stage of development, there was no interaction among the independent loci, and the respective loci together explained most of the variation in the traits. Three independent RFLP loci were associated with canopy width at the V10 stage, of which one was unique to the trait, while the remaining loci (on LGs C2 and F) were in common with the independent loci for plant height. Canopy width had a strong correlation (r=0.87) with plant height at the V10 stage. However, mature plant height, lodging, or seed weight had no phenotypic or QTL association with early plant height or canopy width. Received: 10 May 1998 / Accepted: 13 July 1998     

A soybean mapping population specific to the early soybean production system.

The objective of this research was to create a soybean [Glycine max (L.) Merr] genetic resource in the form of a publicly available, well-characterized mapping population specific to maturity groups (MG) used in the early soybean production system. A total of 568 simple sequence repeat (SSR) markers were tested for polymorphism between soybean breeding line DS97-84-1 (MG IV) and germplasm line DT97-4290 (MG IV). A 90-genotype subset of an F2 population from a cross between these lines was evaluated for genetic linkage using 162 polymorphic SSRs, plant height, pod color (L2/l2), flower color (W1/w1) and stem termination (Dt1/dt1). A 1514 cM (Kosambi) genetic map covering 65% of the soybean genome based on 157 linked SSR markers was created. Comparison with the composite soybean genetic map was used to verify map order. Loci for pod color, flower color and stem termination fell in the expected position on the map indicating this is a normally segregating mapping population. Loci for height were identified on linkage groups C2, D1a, D1b, H, L, M and O. MG IV and V soybean genotypes are critical for the early soybean production system widely used in the midsouthern US. However, only two mapping populations have been reported in Soybase for MG IV and V genotypes. Additionally, the parents used in this cross are known to differ in their response to soybean cyst nematode and charcoal rot, which constitute two major pathology threats to Midsouth soybean production. The population and map reported herein represent an important genetic resource for the early soybean production system.     

Genetic mapping of Sorghum bicolor (L.) Moench QTLs that control variation in tillering and other morphological characters

Grain yield of Sorghum bicolor (L.) Moench is significantly influenced by genetically controlled variation in the number of tillers, plant height, time of anthesis, and various other morphological and physiological characters. In this study, a minimum of 27 unique QTLs that control variation in nine morphological traits, including the presence versus the absence and the height of basal tillers, were mapped, and the percentage of additive genetic variance explained by the QTLs was determined in a population of 137 recombinant inbred lines in two environments. Four QTLs explained from 86.3% to 48.9% (depending upon the environment) of the additive genetic variance in the number of basal tillers with heads, and seven QTLs explained from 85.9% to 47.9% of the additive genetic variance in panicle width. It is unlikely that different alleles were segregating in the mapping population at any of the major dwarfing loci, but five QTLs that explained from 65.8% to 52.0% of the additive genetic variance in main-culm height were mapped. QTLs controlling variation in height of the tallest basal tiller, number of basal tillers per basal-tillered plant, panicle length, leaf angle, maturity, and awn length also were mapped. Three or more QTLs were mapped in linkage groups A, E, G, and I, while none were mapped in linkage groups B and D. Several of the QTLs mapped in this study are likely candidates for marker-assisted selection in breeding programs. Received: 20 September 2000 / Accepted: 26 October 2000     

Construction of a Genetic Linkage Map and Genetic Analysis of Domestication Related Traits in Mungbean (Vigna radiata)

The genetic differences between mungbean and its presumed wild ancestor were analyzed for domestication related traits by QTL mapping. A genetic linkage map of mungbean was constructed using 430 SSR and EST-SSR markers from mungbean and its related species, and all these markers were mapped onto 11 linkage groups spanning a total of 727.6 cM. The present mungbean map is the first map where the number of linkage groups coincided with the haploid chromosome number of mungbean. In total 105 QTLs and genes for 38 domestication related traits were identified. Compared with the situation in other Vigna crops, many linkage groups have played an important role in the domestication of mungbean. In particular the QTLs with high contribution were distributed on seven out of 11 linkage groups. In addition, a large number of QTLs with small contribution were found. The accumulation of many mutations with large and/or small contribution has contributed to the differentiation between wild and cultivated mungbean. The useful QTLs for seed size, pod dehiscence and pod maturity that have not been found in other Asian Vigna species were identified in mungbean, and these QTLs may play the important role as new gene resources for other Asian Vigna species. The results provide the foundation that will be useful for improvement of mungbean and related legumes.     

Impact of epistasis and QTL×environment interaction on the developmental behavior of plant height in rice (Oryza sativa L.)

QTLs with epistatic effects and environmental interaction effects for the developmental behavior of plant height in rice were studied by conventional and conditional methods for quantitative trait loci (QTLs) by mapping with a doubled-haploid population of 123 lines from IR64/Azucena in three environments. The results showed that epistatic effects were important and most epistasis could be detected only by conditional QTL mapping, while most non–epistatic QTLs could be detected by both conventional and conditional methods. Many modificative QTLs showed only epistatic effects without their own additive effects at some stages. QTL×environment (QE) interaction effects were detected more often than QTL main effects for plant-height behavior, which might indicate that gene expression could be greatly affected by the environment. No QTLs had effects during the whole of ontogeny. Conditional QTL mapping might be a valid way to reveal dynamic gene expression for the development of quantitative traits, especially for epistatic effects. Received: 19 May 2000 / Accepted: 27 October 2000