Identification of genomic regions associated with seed dormancy in white-grained wheat

Pre-harvest sprouting (PHS) in developing wheat (Triticum aestivum L.) spikes is stimulated by cool and wet weather and leads to a decline in grain quality. A low level of harvest-time seed dormancy is a major factor for PHS, which generally is a larger problem in white-grained as compared to red-grained wheat. We have in this study analyzed seed dormancy levels at the 92nd Zadok growth stage of spike development in a doubled-haploid (DH) white wheat population and associated variation for the trait with regions on the wheat genome. The phenotypic data was generated by growing the parent lines Argent (non-dormant) and W98616 (dormant) and 151 lines of the DH population in the field during 2002 and 2003, at two locations each year, followed by assessment of harvest-time seed dormancy by germination tests. A genetic map of 2681 cM was constructed for the population upon genotyping 90 DH lines using 361 SSR, 292 AFLP, 252 DArT and 10 EST markers. Single marker analysis of the 90 genotyped lines associated regions on chromosomes 1A, 2B, 3A, 4A, 5B, 6B, and 7A with seed dormancy in at least two out of the four trials. All seven putative quantitative trait loci (QTLs) were contributed by alleles of the dormant parent, W98616. The strongest QTLs positioned on chromosomes 1A, 3A, 4A and 7A were confirmed by interval mapping and markers at these loci have potential use in marker-assisted selection of PHS resistant white-grained wheat.     

Detection of QTLs for traits associated with pre-harvest sprouting resistance in bread wheat (Triticum aestivum L.)

Pre-harvest sprouting (PHS) is one of the serious problems for wheat production, especially in rainy regions. Although seed dormancy is the most critical trait for PHS resistance, the control of heading time should also be considered to prevent seed maturation during unfavorable conditions. In addition, awning is known to enhance water absorption by the spike, causing PHS. In this study, we conducted QTL analysis for three PHS resistant related traits, seed dormancy, heading time and awn length, by using recombinant inbred lines from ‘Zenkouji-komugi’ (high PHS resistance) × ‘Chinese Spring’ (weak PHS resistance). QTLs for seed dormancy were detected on chromosomes 1B (QDor-1B) and 4A (QDor-4A), in addition to a QTL on chromosome 3A, which was recently cloned as TaMFT-3A. In addition, the accumulation of the QTLs and their epistatic interactions contributed significantly to a higher level of dormancy. QDor-4A is co-located with the Hooded locus for awn development. Furthermore, an effective QTL, which confers early heading by the Zenkouji-komugi allele, was detected on the short arm of chromosome 7B, where the Vrn-B3 locus is located. Understanding the genetic architecture of traits associated with PHS resistance will facilitate the marker assisted selection to breed new varieties with higher PHS resistance.     

Integration of marker-assisted selection for resistance to pre-harvest sprouting with selection for grain-filling rate in breeding of white-kernelled wheat for the Chinese environment

Few Chinese high yielding white-grained wheat cultivars possess sufficient dormancy to avoid pre-harvest sprouting (PHS). Because the field evaluation of PHS is difficult, the identification of informative molecular markers is a priority for improving the level of dormancy. In this report, the effectiveness of phenotypic and genotypic selection was compared. Four microsatellite loci Xbarc57, Xbarc294, Xbarc310 and Xbarc321, mapped on the short arm of chromosome 3A, were used for selection in white-grained wheat F₅ lines which were also selected on the basis of their grain filling rate (GFR). One of these (later designated cv. Zhongmai911) was further selected on the basis of its allelic constitution at the four SSR loci. This cultivar combines a high level of PHS resistance with high grain yield. The results suggested that rapid GFR and PHS resistance can be bred simultaneously.     

Comparative analysis of Australian and Canadian barleys for seed dormancy and malting quality

The association between high malting quality and pre-harvest sprouting (PHS) susceptibility is a key challenge when developing new malting barley varieties. A new malting barley variety Baudin has successfully combined high malting quality and PHS tolerance. A doubled haploid population was developed for mapping PHS tolerance and seed dormancy from a cross of Baudin?×?AC Metcalfe using 233 molecular markers. Three QTLs were mapped for seed dormancy based on the standard germination test at 24, 48 and 72?h. One major QTL was mapped to the long arm of chromosome 5H controlling seed dormancy and PHS tolerance from Baudin. Two other minor QTLs were identified from Baudin on chromosomes 3 and 7H. QTL/QTL interaction was detected for seed dormancy between chromosomes 3 and 5H. The PHS tolerance allele of the 5H QTL from Baudin contributes to higher malt yield without significant impact on diastatic power, beta-glucan content and wort viscosity. QTL from Baudin provide new sources to integrate PHS tolerance and high malting quality.     

Grain dormancy in fixed lines of white-grained wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.) grown under controlled environmental conditions

Pre-harvest sprouting (PHS) in wheat (Triticum aestivum L.) can be a significant problem, causing deleterious effects on grain quality. However, the adverse impacts of PHS can be reduced by introgressing genes controlling grain dormancy into white-grained bread wheat. Screening for grain dormancy typically involves germination testing of harvest-ripe grain grown in a glasshouse or field. However, the more uniform environmental conditions provided by temperature controlled glasshouses (i.e. controlled environmental conditions—CEC) may provide significant benefits for the assessment of grain dormancy. In this study, the dormancy phenotype of grain grown under CEC incorporating an extended photoperiod, was compared with 2 years of data from field grown material. Four dormant double haploid lines (derived from SW95-50213 and AUS1408) and two locally adapted non-dormant cultivars EGA Gregory and EGA Wills were compared in three replicated experiments grown under CEC (22 ± 3°C and 24 h photoperiod). The germination response of harvest-ripe grain was examined to assess the expression of grain dormancy. Two measures of germination, the predicted time to 50% germination (G ₅₀) and a weighted germination index, both clearly differentiated dormant and non-dormant lines grown under CEC. In addition, levels of grain dormancy were similar to field-grown plants. These results demonstrated that CEC with an extended photoperiod can be used for rapid and reliable characterisation of grain dormancy in fixed lines of bread wheat.     

Mapping and validation of PCR-based markers associated with a major QTL for seed dormancy in wheat

Seed dormancy is one of the important factors controlling pre-harvest sprouting (PHS) resistance in wheat. We identified a major quantitative trait locus (QTL) for seed dormancy on the long arm of wheat chromosome 4A (4AL) via simple sequence repeat (SSR)-based genetic mapping using doubled haploid lines from a cross between Japanese PHS resistant variety ‘Kitamoe’ and the Alpine non-resistant variety “Münstertaler” (K/M). The QTL explained 43.3% of total phenotypic variation for seed dormancy under greenhouse conditions. SSR markers flanking the QTL were assigned to the chromosome long arm fraction length 0.59–0.66 on the basis of chromosome deletion analysis, suggesting that the gene(s) controlling seed dormancy are probably located within this region. Under greenhouse conditions, the QTL explained 28.5 and 39.0% of total phenotypic variation for seed dormancy in Haruyutaka/Leader (HT/L) and OS21-5/Haruyokoi (O/HK) populations, respectively. However, in field conditions, the effect was relatively low or not significant in both the K/M and HT/L populations. These markers were considered to be widely useful in common with various genetic backgrounds for improvement of seed dormancy through the use of marker-assisted selection. Further detailed research using near isogenic lines will be needed to define how this major QTL interacts with environmental conditions in our area.     

Screening for grain dormancy in segregating generations of dormant × non-dormant crosses in white-grained wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Pre-harvest sprouting (PHS) in wheat (Triticum aestivum L.) is a significant problem. Introgression of genes controlling grain dormancy into white-grained bread wheat is one means of improving resistance to PHS. In this study seven dormant (containing the SW95-50213 and AUS1408 sources) × non-dormant crosses were produced to investigate the effectiveness of selection for grain dormancy in early segregating generations. Each generation (F₁–F₄) was grown in a temperature controlled glasshouse with an extended photoperiod (i.e. continuous light). F₂ and F₃ generations were subject to selection. Five hundred harvest-ripe grains were tested for germination over a 14 day period, and the 100 most dormant grains were retained and grown-on to produce the next generation within each cross. The response to selection was assessed through analysis of the time to 50% germination (G₅₀) in the F₂, F₃ and F₄ generations. In addition, changes in marker class frequencies for two SSR markers (barc170 and gpw2279) flanking a known quantitative trait locus (QTL) for grain dormancy on chromosome 4A were assessed in DNA from F₂ plants selected from early germinating (non-dormant) and late germinating (dormant) phenotypic extremes within each cross. Selection for grain dormancy in the F₂ and F₃ generations effectively recovered the dormant phenotype in all seven crosses, i.e. the F₄ generation was not significantly different from the dormant parent. Further, selection based on individual F₂ grains changed marker class frequencies for the 4A dormancy QTL; in most cases eliminating the marker class homozygous for the non-dormant alleles. Application of this screening method will enable breeders to better select for grain dormancy and may lead to development of new cultivars offering effective resistance to PHS in the near future.     

Mapping quantitative trait loci controlling pre-harvest sprouting resistance in a red × white seeded spring wheat cross

Hard white wheat (Triticum aestivum L.) is a value-added product because of its processing advantages over red wheat; however, white wheat tends to be more susceptible to pre-harvest sprouting (PHS). To identify quantitative trait loci (QTLs) associated with PHS tolerance, we developed a doubled haploid (DH) mapping population from the cross AC Domain (red seeded) × White-RL4137 (white seeded). A genetic map was constructed using microsatellite markers located on chromosome groups 3, 4, 5 and 6. A population of 174 DH lines was characterized for important aspects of PHS including sprouting index, germination index, Hagberg falling number and seed coat colour. A total of 11 QTLs were identified on group 3 chromosomes and on chromosome 5D. Seven QTLs associated with the PHS traits were found to be co-incident with seed coat colour on chromosomes 3A, 3B and 3D. The 5D PHS QTL was notable because it is independent of seed coat colour.     

QTL-based analysis of heterosis for number of grains per spike in wheat using DH and immortalized F 2 populations

To map quantitative trait loci (QTL) and heterotic loci (HL) related to grain number per spike (GNS), 168 double haploid (DH) populations derived from Huapei?3?×?Yumai?57 and an immortalized F ₂ population (IF ₂) generated by randomly permutated intermating of these DH populations were investigated. Using inclusive composite interval mapping (ICIM), a total of nine and eight significant QTLs for GNS were detected in three different environments in DH and IF ₂ populations, respectively. QTLs on chromosomes?1A, 2B, 3B, and 6A were observed between two populations. Five QTLs were detected on chromosome?1A. Of these QTLs, QGns1A-1 was a major QTL explaining 31.25?% of phenotypic variation. QGns2B-2 detected on chromosome?2B had the most significant additive effects, explaining 46.75?% of phenotypic variation with the favorable allele contributed by Yumai?57 corresponding to an increase of 5.69?kernels. Mid-parent heterosis of each cross in the IF ₂ population was used to map heterotic quantitative trait loci. A total of 17 HLs were detected. QTLs and HLs on chromosomes?2B and 6A were observed in the IF ₂ population. Three HLs, QHgns1B-2, QHgns2B, and QHgns6A-1, were detected in two environments and expressed stably. These results showed that some intervals on chromosomes?1B, 2B, and 6A play an important role in GNS heterosis in wheat, improving understanding of this phenomenon.     

Mapping QTLs for pre-harvest sprouting traits in the spring wheat cross ‘RL4452/AC Domain’

Pre-harvest sprouting (PHS) in spring wheat (Triticum aestivum L.) is a major downgrading factors for grain producers and can significantly reduce end-use quality. PHS resistance is a complex trait influenced by genotype, environment and plant morphological factors. A population of 185 doubled haploid (DH) lines from the spring wheat cross ‘RL4452/AC Domain’ were used as the mapping population to detect quantitative trait loci (QTLs) associated with three PHS traits, germination index (GI), sprouting index (SI) and falling number (FN). Six major QTLs linked with PHS traits were mapped on bread wheat chromosomes 3A, 3D, 4A (2 loci), 4B and 7D. ‘AC Domain’ alleles contributed to PHS resistance on 3A, 4A (locus-2) and 4B, and ‘RL4452’ alleles contributed resistance on 3D, 4A (locus-1) and 7D. QTLs detected on chromosome 4B controlling FN (QFN.crc-4B), GI (QGI.crc-4B) and SI (QSI.crc-4B) were coincident, and explained the largest amount of phenotypic variation in FN (22%), GI (67%) and SI (26%), respectively.     

QTL analysis for resistance to preharvest sprouting in rice (Oryza sativa)

Preharvest sprouting (PHS) is caused by early breaking of seed dormancy. In Sichuan, a major hybrid rice seed production area of China, PHS in hybrid seeds originated from ‘G46A’ parent may lead to severe yield loss, causing serious damage to agricultural production. To detect quantitative trait loci (QTLs) governing PHS, we developed an F₂ population of 164 plants derived from ‘G46B’ and ‘K81’, a near‐isogenic introgression line of G46B, with high level of resistance to PHS. PHS was evaluated under controlled field and laboratory conditions. Using simple sequence repeat markers, we constructed a linkage map from this population and identified three QTLs for PHS, namely qPSR2, qPSR5 and qPSR8, which were located on chromosomes 2, 5 and 8, respectively. Among these QTLs, qPSR8, residing in the interval between RM447 and RM3754 on chromosome 8, was the major QTL controlling PHS, for it had a relative high logarithm of the odds (LOD) score and explained 43.04% of the phenotypic variation. These results were correspondent to those identified in extreme low germination rate plants (ELGP) using linkage and linkage disequilibrium. At all loci, ‘K81’ was responsible for enhancing the resistance to PHS.     

Quantitative trait loci for grain yield and yield components in a cross between a six-rowed and a two-rowed barley

Summary The positions of quantitative trait loci (QTL) for yield and yield components were estimated using a 85-point linkage map and phenotype data from a F₁-derived doubled haploid (DH) population of barley. Yield and its components were recorded in two growing seasons. Highly significant QTL effects were found for all traits at several sites in the genome. A major portion of the QTL was found on chromosome 2. The effect of the alleles in locus v on thousand grain weight and kernels per ear explained 70–80% of the genetic variation in the traits. QTL × year interaction was found for grain yield. Several different QTL were found within the two-rowed DH lines compared to those found in the six-rowed DH lines. Epistasis between locus v and several loci for yield and yield components indicates that genes are expressed differently in the two ear types. This may explain the difficulties of selecting high yielding lines from crosses between two-rowed and six-rowed barley.Abbreviations DH doubled haploid - QTL quantitative trait locus/loci - RAPD random amplified polymorphic DNA - RFLP restriction fragment length polymorphism - T. Prentice Tystofte Prentice - V. Gold Vogelsanger Gold     

Detection of chromosomes carrying genes for seed dormancy of wheat using the backcross reciprocal monosomic method

Preharvest sprouting (PHS) in wheat represents a major constraint to the production of high‐quality grain. Genetic variation for tolerance to PHS is associated with seed dormancy. The present study was initiated to detect homologous chromosome variation associated with seed dormancy genes in a PHS tolerant cultivar ‘Zenkojikomugi’ (Zen) and ‘Chinese Spring’ (CS) using the backcross reciprocal monosomic method. The most striking effect on variation in seed dormancy was associated with chromosome 3A, and followed by group 4 chromosomes. These chromosomes of Zen increased dormancy compared with the respective CS homologues. Chromosomes 2D and 7D of Zen, and 6A of CS seemed to contribute smaller positive effects on dormancy. Chromosomes 2A of CS and Zen, as well as 3B, were equally effective in enhancing dormancy. The chromosome 3 A factors were independent of the grain‐colour gene R‐A1 because Zen was found to carry the white‐grained R‐A1a allele, which was identical to that of CS, and there was no difference in dormancy between the reciprocal F₁ seeds.     

Enhancing the identification of genetic loci and transgressive segregants for preharvest sprouting resistance in a durum wheat population

Preharvest sprouting reduces grain quality and lowers grade. Characterization of preharvest sprouting resistance is important in selection in breeding for transgressive segregation and understanding the genetics of the trait for identifying QTL. Methods of measuring dormancy and other factors contributing to preharvest sprouting resistance are varied. The objective of this study was to demonstrate the requirement of multiple methods of measurement over multiple durations of germination to maximize understanding of transgressive segregation and QTL for preharvest sprouting resistance within a segregating durum wheat population grown in multiple environments. Ninety-eight durum wheat (Triticum turgidum L. var. durum) recombinant inbred lines (RIL) from a cross of a minimally dormant line, Sentry, by a moderately dormant line, Kyle, and controls were grown in replicated field tests in 1996, 1997 and 1998 and in a growth chamber trial in 1998. Preharvest sprouting was measured from intact spikes as sprouting index or from hand threshed grain as germination index (GI), germination resistance (GR), and percent germination (PG). The threshed grain measures were evaluated using counts at 7, 14 and 21 days intervals from the start of germination. Correlations performed on the measure type and duration using lines within the RIL population showed some discontinuity across environments, type of measure and duration of measure, with counts at extended intervals for PG producing the lowest correlations. The number of transgressive segregant lines varied with environment, duration and type of measure. Different QTL were identified by different types of measures and duration of counts. GI calculated for 7, 14 and 21 days germination count intervals and GR calculated for 21 days identified a highly significant QTL on chromosome1A (QPhsd.spa.-1A.1). GR calculated for 7 days identified a highly significant QTL on 2A (QPhsd.spa.-2A.1) in two different environments, and GI calculated for 21 days and PG at 7 days identified the same highly significant QTL on chromosome 7B (QPhsd.spa.-7B.1). The results indicated that multiple measures and durations of measure intervals must be applied to results collected across different environments to maximize the identification of QTL and transgressive segregants of the population segregating for preharvest sprouting resistance.     

Characterization of DFR allelic variations and their associations with pre-harvest sprouting resistance in a set of red-grained Chinese wheat germplasm

Pre-harvest sprouting (PHS) of wheat greatly reduces the quality and economic value of grain, and PHS resistance is one of the most important traits in wheat breeding. Red-grained wheat varieties are generally more resistant to PHS than white-grained ones; however, some are still susceptible. The red pigment of red-grained wheat is synthesized through the flavonoid biosynthetic pathway, in which the dihydroflavonol-4-reductase gene (DFR) is one of the genes involved in anthocyanin synthesis. In this study, a set of 120 red-grained Chinese wheat cultivars and lines with distinct PHS resistance were used to characterize TaDFR genotype variations and their association with PHS resistance. Whereas no variation or functional variation of TaDFR genes was detected on chromosomes 3A and 3D, a novel TaDFR allele, designated TaDFR-Bb, was explored on chromosome 3B. Compared with TaDFR-Ba, an 8 bp insertion (CTCTAGGA) was identified in the promoter region of TaDFR-B in most of the PHS resistant red-grained wheat varieties and advanced lines. Based on this, a CAPS marker was designed and validated with a set of Chinese red-grained wheat cultivars and lines with distinct PHS resistance. In most cases, TaDFR-Bb was associated with higher PHS resistance. An association study indicated that wheat varieties with the 8 bp insertion (average seed germination index 23.6 %) were significantly more resistant (P < 0.01) to PHS than those without the insertion (average seed germination index 69.5 %). Further study on gene expression demonstrated that the insertion led to increased TaDFR-B expression in cultivars with PHS resistance. Transient expression of TaDFR-B in coleoptiles of wheat cv. Chinese Spring revealed that increasing TaDFR gene expression did not induce the synthesis of anthocyanins.     

Genetic analysis of pre-harvest sprouting in a durum wheat cross

Pre-harvest sprouting of durum wheat (Triticum turgidum L. var durum) reduces commercial grade, although the actual effects on processing quality are controversial. Little is known about the genetics of the dormancy component of pre-harvest sprouting resistance in durum. We studied the segregation of dormancy in 98 recombinant inbred lines from a cross of a relatively non-dormant line, CI13102, with a moderately dormant line, Kyle. The lines and parents were grown in field tests over three years, 1996, 1997 and 1998. Spikes were collected at approximately 20% moisture and stored at −23 ∘C. Hand-threshed grain of the lines was germinated, and number of seeds germinated was counted each day. A germination resistance index was calculated to characterize dormancy. Dormancy appeared to be complexly inherited in this cross. Lines were observed that were significantly (P < 0.05) more dormant than the parents. The lines transgressive for dormancy expressed in different combinations of the three environments, indicating an environmental interaction. DNA of lines and parents was tested with simple sequence repeat primers and AFLPs that were used in quantitative trait loci (QTL) analysis of dormancy. Significant QTLs for dormancy were found, with the most notable being on chromosome 1A, where other QTLs for pre-harvest sprouting resistance have been reported in common wheat.     

Conferring resistance to pre-harvest sprouting in durum wheat by a QTL identified in <Emphasis Type="Italic">Triticum spelta</Emphasis>

Pre-harvest sprouting (PHS) causes significant yield loss and degrade the end-use quality of wheat, especially in regions with prolonged wet weather during the harvesting season. Unfortunately, the gene pool of Triticum durum (tetraploid durum wheat) has narrow genetic base for PHS resistance. Therefore, finding out new genetic resources from other wheat species to develop PHS resistance in durum wheat is of importance. A major PHS resistance QTL, Qphs.sicau-3B.1, was mapped on chromosome 3BL in a recombinant inbred line population derived from ‘CSCR6’ (Triticum spelta), a PHS resistant hexaploid wheat and ‘Lang’, a PHS susceptible Australian hexaploid wheat cultivar. This QTL, Qphs.sicau-3B.1, is positioned between DArT marker wPt-3107 and wPt-6785. Two SCAR markers (Ph3B.1 and Ph3B.2) were developed to track this major QTL and were used to assay a BC₂F₈ tetraploid population derived from a cross between the durum wheat ‘Bellaroi’ (PHS susceptible) and ‘CSCR6’ (PHS resistant). Phenotypic assay and marker-assisted selection revealed five stable tetraploid lines were highly PHS resistant. This study has successfully established that PHS-resistance QTL from hexaploid wheat could be efficiently introgressed into tetraploid durum wheat. This tetraploid wheat germplasm could be useful in developing PHS resistant durum cultivars with higher yield and good end-use quality.     

Mapping quantitative trait loci for yield-related traits in Chinese cabbage (Brassica rapa L. ssp. pekinensis)

Chinese cabbage (Brassica rapa L. ssp. pekinensis) is one of the most important vegetables in China. However, the inheritance of yield-related traits in Chinese cabbage is poorly understood to date. To map quantitative trait loci (QTL) for yield-related traits in Chinese cabbage, a genetic linkage map was constructed with 192 doubled haploid (DH) lines. The genetic map was constructed based on 190 sequence-related amplified polymorphisms and 43 simple sequence repeats. QTL mapping was conducted for 11 yield-related traits in 170 DH lines derived from a cross between two diverse Chinese cabbage lines, ‘WZ’ and ‘FT’, under different environmental conditions. A total of 46 main QTL (M-QTL) and 7 epistatic QTL (E-QTL) were identified. The phenotypic variation explained by each M-QTL and E-QTL ranged from 4.85 to 25.06 % and 1.85 to 13.29 %, respectively. The QTL-by-environment interactions were detected using the QTLNetwork 2.0 program in joint analyses of multi-environment phenotypic values. The phenotypic variation explained by each QTL and by QTL × environment interaction was 1.14–4.24 % and 0.00–1.26 %, respectively. Our results provide a better understanding of the genetic factors controlling leaf and head-related traits in Chinese cabbage.     

Increased grain dormancy in white-grained wheat by introgression of preharvest sprouting tolerance QTLs

White-grained wheat cultivars have long been recognized to be less resistant to preharvest sprouting (PHS) than the red-grained ones. Previously two QTLs for grain dormancy, QPhs.ocs-3A.1 (QPhs-3AS) and QPhs.ocs-4A.1 (QPhs-4AL) were identified in a highly dormant Japanese red wheat, Zenkoujikomugi (Zen). Aiming at improvement of PHS tolerance in white-grained wheat, the introgression effect of these two QTLs in a white-grained population consisting of 40 recombinant inbred lines (RILs) developed from a cross between Zen and white-grained Spica was examined here. Random 20 RILs with red grains were also developed from the same cross and used as a control population. The RILs were grown in the field and in the glasshouse to evaluate the grain dormancy by germination test. Several SSR markers closely linked to the QPhs-3AS and QPhs-4AL were used to estimate the alleles at the QTLs. Dormancy variation in the RILs was significantly associated with the differences for grain color and the alleles at QPhs-3AS over several years. Although allelic variation was detected in a SSR marker closely linked to QPhs-4AL there was no difference in germination data between the Zen-allele and the Spica-allele groups. As expected, the red-grained RILs with the Zen allele at QPhs-3AS were the most dormant. Some white-grained RILs with the Zen allele at QPhs-3AS showed higher dormancy compared to the red-grained RILs with the alternative allele. These results demonstrated that introgression of the QPhs-3AS gene could contribute to the increased grain dormancy in white-grained wheat.     

Molecular and physical mapping of genes affecting awning in wheat

Quantitative trait loci (QTL) for three traits related to awning (awn length at the base, the middle and the top of the ear) in wheat were mapped in a doubled‐haploid line (DH) population derived from the cross between the cultivars ‘Courtot’ (awned) and ‘Chinese Spring’ (awnless) and grown in Clermont‐Ferrand, France, under natural field conditions. A molecular marker linkage map of this cross that was previously constructed based on 187 DH lines and 550 markers was used for the QTL mapping. The genome was well covered (more than 95%) and a set of anchor loci regularly spaced (one marker every 20.8 cM) was chosen for marker regression analysis. For each trait, only two consistent QTL were identified with individual effects ranging from 8.5 to 45.9% of the total phenotypic variation. These two QTL cosegregated with the genes Hd on chromosome 4A and B2 on chromosome 6B, which are known to inhibit awning. The results were confirmed using ‘Chinese Spring’ deletion lines of these two chromosomes, which have awned spikes, while ‘Chinese Spring’ is usually awnless. No quantitative trait locus was detected on chromosome 5A where the B1 awn‐inhibitor gene is located, suggesting that both ‘Courtot’ and ‘Chinese Spring’ have the same allelic constitution at this locus. The occurrence of awned speltoid spikes on the deletion lines of this chromosome suggests that ‘Chinese Spring’ and ‘Courtot’ have the dominant B1 allele, indicating that B1 alone has insufficient effect to induce complete awn inhibition.