QTL analysis of resistance to Fusarium head blight in wheat using a 'Frontana'-derived population

Fusarium head blight (FHB or head scab) has become a major limiting factor for sustainable wheat (Triticum aestivum L.) production around the world. For quantitative trait loci (QTL) analysis of resistance to FHB, F₃ plants and F_{3 : 5} lines, derived from a ‘Frontana’ (moderately resistant)/‘Seri82’ (susceptible) cross, were spray‐inoculated in 2001 and 2002, respectively. Artificial inoculations were carried out under field conditions. Of 273 SSR and AFLP markers, 250 could be mapped and they yielded 42 linkage groups, covering a genetic distance of 1931 cM. QTL analysis was based on the constructed linkage map and area under the disease progress curve (AUDPC). The analyses revealed three consistent QTLs associated with FHB resistance on chromosomes 1BL, 3AL and 7AS explaining 7.9%, 7.7% and 7.6% of the phenotypic variation, respectively, above 2 years. The results confirmed the previously described resistance QTL of ‘Frontana’ on chromosome 3AL. A combination of ‘Frontana’ resistance with ‘Sumai‐3’ resistance may lead to lines with augmented resistance expression.     

QTLs for Fusarium head blight response in a wheat DH population of Wangshuibai/Alondra‘s’

Summary A doubled haploid (DH) wheat population derived from the cross Wangshuibai/Alondra‘s’ was developed through chromosome doubling of haploids generated by anther culture of hybrids. Fusarium head blight (FHB) was evaluated for three years from 2001 to 2003 in Jianyang, Fujian Province, China, where epidemics of FHB have been consistently severe. After 307 pairs of simple sequence repeat (SSR) primers were screened, 110 pairs were polymorphic between Wangshuibai and Alondra`s’, and used to construct a genetic linkage map for detection of quantitative trait loci (QTLs). A stable QTL for low FHB severity was detected on chromosomes 3B over all three years, and QTLs on chromosomes 5B, 2D, and 7A were detected over two years. Additional QTLs on chromosomes 3A, 3D, 4B, 5A, 5D, 6B and 7B showed marginal significance in only one year. Six QTLs were detected when phenotypic data from three years were combined. In addition, significant additive-by-additive epistasis was detected for a QTL on 6A although its additive effect was not significant. Additive effects (A) and additive-by-additive epistasis (AA) explained a major portion of the phenotypic variation (76.5%) for FHB response. Xgwm533-3B and Xgwm335-5B were the closest markers to QTLs, and have potential to be used as selectable markers for marker-assisted selection (MAS) in wheat breeding programs.     

Validation of a major QTL for scab resistance with SSR markers and use of marker-assisted selection in wheat

The objectives of this study were to validate the major quantitative trait locus (QTL) for scab resistance on the short arm of chromosome 3B in bread wheat and to isolate near‐isogenic lines for this QTL using marker‐assisted selection (MAS). Two resistant by susceptible populations, both using ‘Ning7840’ as the source of resistance, were developed to examine the effect of the 3BS QTL in different genetic backgrounds. Data for scab resistance and simple sequence repeat (SSR) markers linked to the resistance QTL were analyzed in the F_2:3 lines of one population and in the F_3:4 lines of the other. Markers linked to the major QTL on chromosome 3BS in the original mapping population (‘Ning7840’/‘Clark’) were closely associated with scab resistance in both validation populations. Marker‐assisted selection for the QTL with the SSR markers combined with phenotypic selection was more effective than selection based solely on phenotypic evaluation in early generations. Marker‐assisted selection of the major QTL during the seedling stage plus phenotypic selection after flowering effectively identified scab resistant lines in this experiment. Near‐isogenic lines for this 3BS QTL were isolated from the F₆ generation of the cross ‘Ning7840’/‘IL89‐7978’ based on two flanking SSR markers, Xgwm389 and Xbarc147. Based on these results, MAS for the major scab resistance QTL can improve selection efficiency and may facilitate stacking of scab resistance genes from different sources.     

Molecular characterization of <Emphasis Type="Italic">Fusarium</Emphasis> head blight resistance from wheat variety Wangshuibai

Fusarium head blight (FHB) is a destructive disease of wheat worldwide. FHB resistance genes from Sumai 3 and its derivatives such as Ning 7840 have been well characterized through molecular mapping. In this study, resistance genes in Wangshuibai, a Chinese landrace with high and stable FHB resistance, were analyzed through molecular mapping. A population of 104 F₂-derived F₇ recombinant inbred lines (RILs) was developed from the cross between resistant landrace Wangshuibai and susceptible variety Alondras. A total of 32 informative amplified fragment length polymorphism (AFLP) primer pairs (EcoRI/MseI) amplified 410 AFLP markers segregating among the RILs. Among them, 250 markers were mapped in 23 linkage groups covering a genetic distance of 2,430 cM. In addition, 90 simple sequence repeat (SSR) markers were integrated into the AFLP map. Fifteen markers associated with three quantitative trait loci (QTL) for FHB resistance (P < 0.01) were located on two chromosomes. One QTL was mapped on 1B and two others were mapped on 3B. One QTL on 3BS showed a major effect and explained up to 23.8% of the phenotypic variation for type II FHB resistance.     

Phenotypic selection for high resistance to Fusarium head blight after introgression of quantitative trait loci (QTL) from exotic spring wheat and verification by simple sequence repeat markers a posteriori

Fusarium head blight (FHB) has become an important disease of wheat. We introgressed three resistance quantitative trait loci (QTL) alleles on chromosomes 3B, 5A (from CM82036) and 3A (from ‘Frontana’) into European elite spring wheat and performed phenotypic selection among double‐cross (DC) derived progeny in generations DCF₂ and DCF₃. After recombination and selfing, we analysed 135 phenotypically selected progeny by simple sequence repeat (SSR) markers linked to the QTL. In a second experiment, we forwarded the best 20 progeny for a further two generations by pedigree selection. Progeny were inoculated at two to four locations with Fusarium culmorum and the percentage of infected spikelets per plot was estimated. Both experiments show that phenotypic selection was highly effective. One‐hundred out of 135 phenotypically selected DCF_1:3 progeny had the combination of donor‐QTL alleles (3B + 5A + 3A, 3B + 5A) with the highest effects on FHB resistance. In the subsequent generations, sufficient genotypic variance was detected. The best F_5:7 bulks had similar resistance to the donor CM82036. The FHB rating was reduced in total by 45% points compared to the parental mean. QTL with high effects can be detected solely by phenotypic selection after targeted introgression.     

Genetic mapping and inheritance of Russian wheat aphid resistance gene in accession IG 100695

The Russian wheat aphid (RWA), Diuraphis noxia (Kurdjumov), is an important pest of small‐grain cereals, particularly wheat, worldwide. The most efficient strategy against the RWA is to identify sources of resistance and to introduce them into susceptible wheat genotypes. This study was conducted to determine the mode of inheritance of the RWA resistance found in ICARDA accession IG 100695, to identify wheat microsatellite markers closely linked to the gene and to map the chromosomal location of the gene. Simple sequence repeat (SSR) marker scores were identified in a mapping population of 190 F₂ individuals and compared, while phenotypic screening for resistance was performed in F_2 : 3 families derived from a cross between ‘Basribey’ (susceptible) and IG 100695 (resistant). Phenotypic segregation of leaf chlorosis and rolling displayed the effect of a single dominant gene, temporarily denoted Dn100695, in IG 100695. Dn100695 was mapped on the short arm of chromosome 7D with four linked SSR markers, Xgwm44, Xcfd14, Xcfd46 and Xbarc126. Dn100695 and linked SSR markers may be useful for improving resistance for RWA in wheat breeding.     

Amplified fragment length polymorphism- and simple sequence repeat-based molecular tagging and mapping of greenbug resistance gene Gb3 in wheat

The greenbug, Schizaphis graminum (Rondani), is the most economically damaging aphid pest of wheat in the southern Great Plains of the USA. In this study, the single, dominant greenbug resistance gene, Gb3, was molecularly tagged and genetically mapped using amplified fragment length polymorphism (AFLP) and simple sequence repeat(SSR) markers. Three AFLP loci were associated with the Gb3 locus in linkage analysis with 75 F_2:3 families from the cross between two near‐isogenic lines (NILs) for Gb3,‘TXGBE273’ and ‘TXGBE281′. Two of these loci, XMgcc Pagg and Xmagg Patg cosegregate with Gb3 in the population analysed. Further analysis indicated that XMgcc Pagg and Xmagg Patg are specific for the Gb3 locus in diverse genetic backgrounds. Two SSR markers, Xgwm111 and Xgwm428 previously mapped in wheat chromosome 7D, were shown to be linked with Gb3, 22.5 cM and 33.1 cM from Gb3, respectively, in an F₂ population of ‘Largo’בTAM 107’, suggesting that Gb3 is located in the long arm of chromosome 7D. The two AFLP markers cosegregating with Gb3 are valuable tools in developing molecular markers for marker‐assisted selection of greenbug resistance in wheat breeding.     

YM型小麦温敏雄性不育系不育基因的QTL定位

YM型小麦温敏雄性不育系的不育基因被定位在1Bs染色体片段上, 但已发现的相邻分子标记与该基因的遗传距离较大, 达10 cM以上。为寻找与该基因连锁更紧密的分子标记, 以YM型温敏雄性不育系ATM3314与恢复系中国春杂交的F₂代200株为作图群体, 从1Bs的22个SSR引物中筛选出5个在亲本和F₂代中分离的SSR引物, 构建了1个包含5个标记的1Bs局部遗传连锁图谱。结合F₂代个体的育性调查, 采用复合区间作图法在YM型温敏雄性不育系的1Bs染色体上检测到不育基因的1个主效QTLrfv1-1和1个微效QTLrfv1-2。rfv1-1位于SSR标记Xgwm18和Xwmc406之间, 与两标记的遗传距离分别为6.0 cM和4.6 cM, LOD值为8.80, 加性效应23.87, 显性效应10.44, 可解释表型变异的23.91%; rfv1-2位于Xwmc406和Xbarc8之间, 与两标记的遗传距离分别为4.0 cM和3.4 cM, LOD值为3.10, 加性效应17.59, 显性效应5.99, 可解释表型变异的7.78%。本研究初步定位了YM型小麦温敏雄性不育系1Bs染色体片段上不育基因的QTL, 为进一步准确定位该基因奠定了基础。     

Mapping of quantitative trait loci conferring resistance to bacterial wilt in tobacco (Nicotiana tabacum L.)

Tobacco bacterial wilt (TBW) is one of the most serious tobacco diseases in the world. Studies have shown that tobacco resistance to TBW is quantitatively inherited. This study aimed to map quantitative trait loci (QTL) conferring TBW resistance. An F_2 : 3 population containing 237 lines was developed from a cross between two flue‐cured tobacco cultivars, ‘Yanyan 97’ (YY97; moderately resistant to TBW) and ‘Honghua Dajinyuan’ (HD; highly susceptible to TBW), and a linkage map consisting of 201 simple sequence repeats (SSR) markers and spanning a total length of 2326.7 cM was constructed based on the population. Field experiments were conducted 2011 and 2012, and disease symptoms were investigated three times in each year. The phenotypic data were analysed either separately or jointly for QTL mapping using the software QTLNetwork 2.1. Eight QTL with significant main effects were mapped on chromosomes 2, 6, 12, 17 and 24. A major QTL (qBWR17a) was detected on chromosome 17, which explained up to 30% of the phenotypic variation. The results can facilitate marker‐assisted selection (MAS) in TBW resistance breeding programme.     

Identification and confirmation of quantitative trait loci for stachyose content in soybean seed

Stachyose is an unfavorable sugar in soybean meal that causes flatulence for non‐ruminant animals. Understanding the genetic control of stachyose in soybean will facilitate the modification of stachyose content at the molecular level. The objective of this study was to identify quantitative trait loci (QTL) associated with seed stachyose content using simple sequence repeat (SSR) and single nucleotide polymorphism (SNP) markers. A normal stachyose cultivar, ‘Osage’, was crossed with a low stachyose line, V99‐5089, to develop a QTL mapping population. Two parents were screened with 33 SSR and 37 SNP markers randomly distributed on chromosome 10, and 20 SSR and 19 SNP markers surrounding a previously reported stachyose QTL region on chromosome 11. Of these, 5 SSR and 16 SNP markers were used to screen the F_3:4 lines derived from ‘Osage’ x V99‐5089. Seed samples from F_3:5 and F_3:6 lines were analyzed for stachyose content using high‐performance liquid chromatography (HPLC). Composite interval mapping analysis indicated that two stachyose QTL were mapped to chromosome 10 and 11, explaining 11% and 79% of phenotypic variation for stachyose content, respectively. The SSR/SNP markers linked to stachyose QTL could be used in breeding soybean lines with desired stachyose contents. Chi‐square tests further indicated that these two QTL probably represent two independent genes for stachyose content. Therefore, a major QTL was confirmed on chromosome 11 and a novel QTL was found on chromosome 10 for stachyose content.     

Fusarium head blight resistance derived from Lophopyrum elongatum chromosome 7E and its augmentation with Fhb1 in wheat

The objective of this study was to assess the effectiveness of Fusarium head blight (FHB) resistance derived from wheatgrass Lophopyrum elongatum chromosome 7E and to determine whether this resistance can augment resistance in combination with other FHB resistance quantitative trait loci (QTL) or genes in wheat. The ‘Chinese Spring’–Lophopyrum elongatum disomic substitution line 7E(7B) was crossed to three wheat lines: ‘Ning 7840’, L3, and L4. F₂ populations were evaluated for type II resistance with the single‐floret inoculation method in the greenhouse. Simple sequence repeat markers associated with Fhb1 in ‘Ning 7840’ and L4 and markers located on chromosome 7E were genotyped in each population. Marker–trait association was analysed with one‐way or two‐way analysis of variance. The research showed that, in the three populations, the average number of diseased spikelets (NDS) in plants with chromosome 7E is 1.2, 3.1 and 3.2, vs. NDS of 3.3, 7.2 and 9.1 in plants without 7E, a reduction in NDS of 2.1, 4.1 and 5.9 in the respective populations. The QTL on 7E and the Fhb1 gene augment disease resistance when combined. The effect of the QTL on 7E was greater than that on 3BS in this experiment. Data also suggest that the FHB resistance gene derived from L. elongatum is located on the long arm of 7E.     

Detection of downy and powdery mildew resistance QTL in a ‘Regent’ × ‘RedGlobe’ population

One hundred and eighty six F₁ plants from a ‘Regent’ × ‘RedGlobe’ cross were used to generate a partial linkage map with 139 microsatellite markers spanning all 19 chromosomes. Phenotypic scores for downy mildew, taken over two years, confirmed a major resistance QTL (Rpv3) against downy mildew in the interval VVIN16-cjvh to UDV108 on chromosome 18 of ‘Regent’. This locus explained up to 62 % of the phenotypic variance observed. Additionally a putative minor downy mildew resistance locus was observed on chromosome 1 in one season. A major resistance locus against powdery mildew (Ren3) was also identified on chromosome 15 of ‘Regent’ in the interval UDV116 to VChr15CenGen06. This study established the efficacy of and validated the ‘Regent’-derived downy and powdery mildew major resistance genes/QTL under South African conditions. Closely linked SSR markers for marker-assisted selection and gene pyramiding strategies were identified.     

利用永久F2群体定位小麦株高的QTL

为研究小麦株高的遗传机制,利用DH群体构建了一套包含168个杂交组合的小麦永久F₂群体, 并于2007年种植于山东泰安和山东聊城。构建了一套覆盖小麦21条染色体的遗传连锁图谱并利用该图谱的324个SSR标记对小麦株高进行QTL定位研究,使用基于混合线性模型的QTLNetwork 2.0软件进行QTL分析。在永久F₂群体中定位了7个株高QTL,包括4个加性QTL,一个显性QTL,一对上位性QTL,共解释株高变异的20%,其中位于4D染色体的qPh4D,具有最大的遗传效应,贡献率为7.5%;位于2D 染色体显性效应位点qPh2D,可解释1.6%的表型变异;位于5B~6D染色体上位效应位点,可解释1.7%的表型变异。还发现加性效应、显性效应和上位效应对小麦株高的遗传起重要作用,并且基因与环境具有互作效应,结果表明利用永久F₂群体进行QTL定位研究的方法有助于分子标记辅助育种。     

Mapping of the Vrn-B1 gene in Triticum aestivum using microsatellite markers

An F₂ population segregating for the dominant gene Vrn‐B1 was developed from the cross of the substitution line ‘Diamant/'Miro‐novskaya 808 5A’ and the winter wheat cultivar ‘Bezostaya 1′. Microsatellite markers (Xgwm and Xbarc) with known map locations on chromosome 5B of common wheat were used for mapping the gene Vrn‐B1. Polymorphism between parental varieties was observed for 28 out of 34 microsatellite markers (82%). Applying the quantitative trait loci mapping approach, the target gene was mapped on the long arm of chromosome 5B, closely linked to Xgwm408. The map position of Vrn‐B1 suggests that the gene is homoeologous to other vernalization response genes located on the homoeologous group 5 chromosomes of wheat, rye and barley.     

Fine mapping the QTL qFS‐chr.23, using a CSIL

In an earlier advanced‐backcross quantitative trait locus (QTL) analysis of an interspecific cross of Gossypium hirsutum cv. ‘Xinluzhong 36’(‘XLZH36’) and G. barbadense cv. ‘Xinhai 21’(‘XH21’), a QTL for fibre strength in the chromosome segment introgression line IL23‐09 was analysed. Single marker analysis revealed that the markers on chro.23 were associated with fibre strength. Using composite interval mapping with the F₂ population (1296 plants), a QTL for fibre strength was detected on chro. 23. The QTL explained 8.9% and 15.9% of phenotypic variances in the F₂ and F_2 : 3 generations, respectively. Substitution mapping suggested that the QTL was located at a physical distance of 23.4 kb between the markers BNL1414 and the single nucleotide polymorphism (SNP) locus D09_43776813 C‐G. We designated this QTL as qFS‐chr.23 (quantitative trait locus for fibre strength on chro.23). This work provides a valuable genetic resource for the breeding of high fibre quality in cotton and will facilitate future efforts for map‐based cloning.     

A QTL for early heading in wheat cultivar Suwon 92

Heading date is an important trait that determines wheat adaptation to environments. A recombinant inbred line (RIL) population derived from CI 13227 × Suwon 92 was employed to tag the quantitative trait locus (QTL) for early heading in Suwon 92. This population was phenotyped for heading date in 1994, 1995, and 1997, and analyzed with AFLP and SSR markers. Two AFLP markers (XGCTG.CGCT118 and XGCTG.CGCT60) closely associated with heading date were identified. Across years, XGCTG.CGCT118 and XGCTG.CGCT60 explained 40.4% and 32.2% of the total phenotypic variances, respectively. Interval analysis revealed a major QTL for heading date, designated QHd.pser-2DS, between AFLP marker XGCTG.CGCT118 and SSR marker Xgwm261. Based on the linkage map, QHd.pser-2DS was about 41.2 cM proximal to the distal end of chromosome 2DS, and explained 40.5% of the phenotypic variance across three years. The identified markers associated with the early heading QTL have the potential to be used in wheat breeding programs.     

Linkage map construction and mapping QTL for cotton fibre quality using SRAP, SSR and RAPD

Tetraploid cotton is one of the most extensively cultivated species. Two tetraploid species, Gossypium hirsutum L. and G. barbadense L., dominate the world's cotton production. To better understand the genetic basis of cotton fibre traits for the improvement of fibre quality, a genetic linkage map of tetraploid cotton was constructed using sequence‐related amplified polymorphisms (SRAPs), simple sequence repeats (SSRs) and random amplified polymorphic DNAs (RAPDs). A total of 238 SRAP primer combinations, 368 SSR primer pairs and 600 RAPD primers were used to screen polymorphisms between G. hirsutum cv. Handan208 and G. barbadense cv. Pima90 which revealed 749 polymorphic loci in total (205 SSRs, 107 RAPDs and 437 SRAPs). Sixty‐nine F₂ progeny from the interspecific cross of ‘Handan208’בPima90’ were genotyped with the 749 polymorphic markers. A total of 566 loci were assembled into 41 linkage groups with at least three loci in each group. Twenty‐eight linkage groups were assigned to corresponding chromosomes by SSR markers with known chromosome locations. The map covered 5141.8 cM with a mean interlocus space of 9.08 cM. A × test for significance of deviations from the expected ratio (1: 2: 1 or 3: 1) identified 135 loci (18.0%) with skewed segregation, most of which had an excess of maternal parental alleles. In total, 13 QTL associated with fibre traits were detected, among which two QTL were for fibre strength, four for fibre length and seven for micronaire value. These QTL were on nine linkage groups explaining 16.18‐28.92% of the trait variation. Six QTL were located in the A subgenome, six QTL in the D subgenome and one QTL in an unassigned linkage group. There were three QTL for micronaire value clustered on LG1, which would be very useful for improving this trait by molecular marker‐assisted selection.     

Chromosomal location of powdery mildew resistance gene Pm16 in wheat using SSR marker analysis

The use of resistant cultivars is a most economical way to control powdery mildew (Blumeria graminis f.sp. tritici) in wheat (Triticum aestivum L.). Identification of molecular markers closely linked to resistance genes can greatly increase the efficiency of pyramiding resistance genes in wheat cultivars. The objective of this study was to identify molecular markers closely linked lo the powdery mildew resistance gene Pm16. An F₂ population with 156 progeny was produced from the cross‘Chancellor’(susceptible) ב70281’ (resistant), A total of 45 SSR markers on chromosomes 4A and 5B of wheat and 15 SSRs on chromosome 3 of rice was used lo lest the parents, as well as the resistant and susceptible bulks: the resulting polymorphic markers were used to genotype the F₂ progeny. Results indicated that the SSR marker Xgwm159, located on the short arm of chromosome 5B, is closely linked to Pm16 (genetic distance: 5.3 CM). The cytogenetical data presented in an original report, in combination with this molecular analysis, suggests that Pm16 may he located on a translocated 4A.5BS chromosome.     

Effect of 3B, 5A and 3A QTL for Fusarium head blight resistance on agronomic and quality performance of Canadian winter wheat

The objectives of this study were to investigate (i) the correlations between Fusarium head blight (FHB) index, deoxynivalenol (DON) accumulation and percentage of Fusarium‐damaged kernels (FDK) with agronomic and quality traits and (ii) the effect associated with the presence of single QTLs for FHB resistance on agronomic and quality traits in winter wheat. The population was derived from the cross between ‘RCATL33' (FHB resistance derived from ‘Sumai 3’ and ‘Frontana’) and ‘RC Strategy’. Parental lines and recombinant inbred lines (RILs) were genotyped with SSR markers associated with the 3B, 5A and 3A QTLs. The population was planted in FHB‐inoculated nurseries and in agronomy trials. Lines in the 3B QTL class had the lowest FHB index, DON content and FDK level and did not have a significantly lower yield, thousand kernel weight or protein content compared with the lines grouped in other QTL classes (including no QTL class). Marker‐assisted selection of the 3B QTL for FHB resistance into high‐yielding FHB‐susceptible winter wheat is the recommended approach for the development of lines with increased FHB resistance without significant yield and quality penalties.     

Quantitative trait loci mapping of seed colour,hairy leaf,seedling anthocyanin,leaf chlorosis and days to flowering in F2 population of Brassica rapa L.

A diversity arrays technology (DArT) map was constructed to identify quantitative trait loci (QTL) affecting seed colour, hairy leaf, seedling anthocyanin, leaf chlorosis and days to flowering in Brassica rapa using a F₂ population from a cross between two parents with contrasting traits. Two genes with dominant epistatic interaction were responsible for seed colour. One major dominant gene controls the hairy leaf trait. Seedling anthocyanin was controlled by a major single dominant gene. The parents did not exhibit leaf chlorosis; however, 32% F₂ plants showed leaf chlorosis in the population. A distorted segregation was observed for days to flowering in the F₂ population. A linkage map was constructed with 376 DArT markers distributed over 12 linkage groups covering 579.7 cM. The DArT markers were assigned on different chromosomes of B. rapa using B. rapa genome sequences and DArT consensus map of B. napus. Two QTL (RSC1‐2 and RSC12‐56) located on chromosome A8 and chromosome A9 were identified for seed colour, which explained 19.4% and 18.2% of the phenotypic variation, respectively. The seed colour marker located in the ortholog to Arabidopsis thaliana Transparent Testa2 (AtTT2). Two QTL RLH6‐0 and RLH9‐16 were identified for hairy leaf, which explained 31.6% and 20.7% phenotypic variation, respectively. A single QTL (RSAn‐12‐157) on chromosome A7, which explained 12.8% of phenotypic variation was detected for seedling anthocyanin. The seedling anthocyanin marker is found within the A. thaliana Transparent Testa12 (AtTT12) ortholog. A QTL (RLC6‐04) for leaf chlorosis was identified, which explained 55.3% of phenotypic variation. QTL for hairy leaf and leaf chlorosis were located 0–4 cM apart on the same chromosome A1. A single QTL (RDF‐10‐0) for days to flowering was identified, which explained 21.4% phenotypic variation.