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.     

Substitution analysis of drought tolerance in wheat (Triticum aestivum L.)

Chromosome substitution lines of the wheat variety ‘Cappelle Desprez’ into ‘Chinese Spring’ were tested for drought tolerance in growth chambers in the Martonvásár phytotron. Three different moisture regimes were created: E1, fully irrigated control; E2, mid-season water stress (preanthesis); and E3, terminal water-stress during grain filling. Data were analysed to estimate the chromosomal location of the genes controlling relative water-content (RWC), relative water-loss (RWL), drought-susceptibility index (DSI) and phenotypic stability in each substitution line. Simultaneous consideration indicated that most of the genes controlling these characters are located on chromosomes 1A, 5A, 7A,4B, 5B, 1D, 3D and 5D.     

Genetic control of frost tolerance in wheat (Triticum aestivum L.)

Summary The frost tolerance of winter wheat is one component of winter hardiness. If seedlings are frost resistant, it means that they can survive the frost effect without any considerable damage. To study the genetic control of frost tolerance, an artificial freezing test was used. Frost tolerance is controlled by an additive-dominance system. The results of diallel analyses indicate the importance of both additive and non-additive gene action in the inheritance of this character. The dominant genes act in the direction of lower frost tolerance and the recessive genes in the direction of a higher level of frost tolerance. The results of monosomic and substitution analyses show that at least 10 of the 21 pairs of chromosomes are involved in the control of frost tolerance and winter hardiness. Chromosomes 5A and 5D have been implicated most frequently. The geneFr1 (Frost 1) was located on the long arm of chromosome 5A. Crosses between cultivars, chromosome manipulation and the induction of somaclonal variation may be suitable methods for broadening the gene pool for frost tolerance.     

The low down on association mapping in hexaploid wheat (Triticum aestivum L.)

The past few decades have witnessed hundreds of family-based linkage studies mapping for numerous traits but only a limited number of QTLs were actually cloned, tagged, or used for marker-assisted selection. Although providing valuable information, this conventional approach cannot be scaled up to underpin the incredible amount of phenotypic variation in the form of 266, 589 hexaploid wheat accessions maintained in public germplasm collections. Association mapping has recently emerged as an alternative and more powerful mapping approach where a natural population is surveyed to determine marker-trait associations using linkage disequilibrium (LD). After its first application for milling quality in 2006, association mapping studies in hexaploid wheat are being extended to tag yield traits, protein quality, and tolerance to biotic and abiotic stresses. Advances in genotyping technology and statistical approaches greatly accelerated the shift from conventional linkage-based mapping to LD-based association mapping. Association mapping stands out because of simultaneous utilization of a large number of ex situ-conserved natural variation due to historical recombination events accumulated over centuries.     

Growth habit in common wheat (Triticum aestivum L. em. Thell.)

Summary A study of the Vrn genotypes of 642 spring wheats supports the theory that only Vrn1, Vrn2 and Vrn3 exist in Tricticum aestivum. In none of the varieties investigated Vrn4 was present. Seven varieties, which according to literature carry Vrn4, showed to carry Vrn1, Vrn2 and/or Vrn3. Some varieties were mixtures of Vrn-genotypes, which could mislead geneticists in pooled data analysis. Other causes for misinterpretation of the data could be hybrid necrosis, hybrid dwarfness or a wrong determination of plants with a winter habitus. Only Hope was dominant on another Vrn locus. Its haploid Vrn-genotype is Vrn1 vrn2vrn3 Vrn5.     

Mapping QTLs for potassium-deficiency tolerance at the seedling stage in wheat (Triticum aestivum L.)

Potassium (K) deficiency is a major factor limiting crop development and yield. In this study, 20 traits were investigated at the seedling stage using a doubled-haploid population subjected to normal K and K deprivation treatments. Evaluation of phenotypes expressed under the two K supply conditions revealed that K deprivation was able to decrease values of ten measured traits, whereas values of the other ten traits increased. A total of 65 QTLs were detected across all 21 chromosomes, except for chromosomes 2B, 5A, and 7B. Individual QTLs in the two K supply treatments explained between 5.35 and 39.64 % of the phenotypic variation. Nine QTL clusters (C1–C9) involving 34 QTLs under the different K supply treatments were mapped to chromosomes 1A, 1B, 1D, 3B, 4A, 5B, 6A, and 7B. Our results are helpful for understanding the genetic basis of K deficiency in wheat, and provide useful information for genetic improvement of K deficiency in wheat by marker-assisted selection.     

Relationship between anther culture response and aluminium tolerance in wheat (Triticum aestivum L.)

A combination of in vivo and in vitro selection methods were used to increase aluminium tolerance in wheat using wheat x triticale crosses. Both in vivo and in vitro aluminium treatments significantly influenced the anther culture response. In vivo selection at the seedling stage resulted in significantly higher embryo induction. On induction media containing aluminium, the embryoid induction frequency dropped significantly, but there was an increase in the green plant regeneration frequency. In spite of this effect, all doubled haploid (DH) lines were more tolerant to aluminium in seedling tests than the winter wheat parent. The application of in vivo aluminium selection, before the start of anther culture, increased the probability of obtaining DH lines with significantly higher tolerance, compared to the original population. After three selection cycles of the original populations, there was a significant difference in the root regrowth rate of tolerant and sensitive plants. Both sensitive and tolerant plants showed a decrease due to the presence of aluminium in the induction media, with a greater decrease occurring in sensitive plants. Correlation between the rate of root regrowth in the seedling test and the change in embryo induction was positive, but moderate, emphasising the fact that plants with higher root regrowth tended to be more tolerant of the presence of aluminium in the induction medium. This revised version was published online in August 2006 with corrections to the Cover Date.     

Evaluation of fresh pasta-making properties of extra-strong common wheat (Triticum aestivum L.)

The relationship between characterictics of flour of common wheat varieties and fresh pasta-making qualitites was examined, and the fresh pasta-making properties of extra-strong varieties that have extra-strong dough were evaluated. There was a positive correlation between mixing time (PT) and hardness of boiled pasta, indicating that the hardness of boiled pasta was affected by dough properties. Boiled pasta made from extra-strong varieties, Yumechikara, Hokkai 262 and Hokkai 259, was harder than that from other varieties and commercial flour. There was a negative correlation between flour protein content and brightness of boiled pasta. The colors of boiled pasta made from Yumechikara and Hokkai 262 grown under the condition of standard manuring culture were superior to those of boiled pasta made from other varieties. Discoloration of boiled pasta made from Yumechikara grown under the condition of heavy manuring culture was caused by increase of flour protein content. On the other hand, discoloration of boiled pasta made from Hokkai 262 grown under the condition of heavy manuring culture was less than that of boiled pasta made from Yumechikara. These results indicate that pasta made from extra-strong wheat varieties has good hardness and that Hokkai 262 has extraordinary fresh pasta-making properties.     

Flooding tolerance of spelt (Triticum spelta L.) compared to wheat (Triticum aestivum L.) – A physiological and genetic approach

In marginal, agroclimatic zones, yield is often affected by flooding, but the effect is much less for winter spelt (Triticum spelta L.) than for winter wheat (Triticum aestivum L.). This study evaluates the reaction of a wheat x spelt population (F₅ RILs of Forno x Oberkulmer) to flooding stress in the early phase of germination. Lines with greater tolerance to 48 h flooding just after imbibition showed less electrolyte leakage (r = -0.79) indicating greater membrane integrity and better survival. Five QTL explaining 40.6% of the phenotypic variance for survival to flooding were found, and localized on the chromosomes 2B, 3B,5A, and 7S. The tolerance to 48 h flooding four days after sowing was best correlated with the mean germination time (r = 0.8), indicating that the plants with a fast coleoptile growth during flooding are less susceptible to flooding. Ten QTL were found for seedling growth index after flooding explaining 35.5% of the phenotypic variance. They were localized on chromosomes 2A, 2B, 2D, 3A, 4B, 5A, 5B, 6A, and 7S. Standard varieties of spelt and wheat showed the same tolerance characteristics. The possibility to use marker assisted selection for flooding tolerance is discussed. This revised version was published online in August 2006 with corrections to the Cover Date.     

Relationship between common wheat (Triticum aestivum L.) gluten proteins and dough rheological properties

This paper reports the correlation between the rheological properties of bread wheat dough and the types and quantities of endosperm proteins in 28 common wheat cultivars. Different methods were used to analyse the allelic composition of these cultivars and the relative quantities of the different proteins contributing to the gluten structure. Neither dough strength (W) nor tenacity/extensibility (P/L) correlated with allelic composition. Different wheats with the same allelic composition (i.e., with respect to glutenins) showed different rheological properties. The glutenins were the most influential components of W and P/L, especially the high molecular weight (HMW) glutenin subunits and in particular the type x form. These proteins seem to increase W and are the main constituents of the gluten network. The gliadins and low molecular weight (LMW) glutenin subunits appear to act as a “solvent”, and thus modify the rheological properties of the dough by either interfering with the polymerisation of the HMW glutenin subunits, or by altering the relative amounts of the different types of glutenin available. Thus, the protein subunits coded for by the alleles Glu-B1x7 and Glu-D1x5 stabilised the gluten network, whereas those coded for by Glu-B1x17 and Glu-D1x2 had the opposite effect. Dough properties therefore appear to depend on the glutenin/gliadins balance, and on the ratio of the type x and type y HMW proteins. The influence of external factors seems to depend on the allelic composition of each cultivar.     

Genome-wide QTL analysis for pre-harvest sprouting tolerance in bread wheat

A framework linkage map comprising 214 molecular marker (SSR, AFLP, SAMPL) loci was prepared using an intervarietal recombinant inbred line (RIL) mapping population of bread wheat. The RIL population that was developed from the cross SPR8198 (red-grained and PHS tolerant genotype) × HD2329 (white-grained and PHS susceptible genotype) following single seed descent segregated for pre-harvest sprouting (PHS). The RIL population and parental genotypes were evaluated in six different environments and the data on PHS were collected. Using the linkage map and PHS data, genome-wide single-locus and two-locus QTL analyses were conducted for PHS tolerance (PHST). Single-locus analysis following composite interval mapping (CIM) detected a total of seven QTL, located on specific arms of five different chromosome (1AS, 2AL, 2DL, 3AL and 3BL). These seven QTL included two major QTL one each on 2AL and 3AL. Two of these seven QTL were also detected following two-locus analysis, which resolved a total of four main-effect QTL (M-QTL), and 12 epistatic QTL (E-QTL), the latter involved in 7 QTL × QTL interactions. Interestingly, none of these M-QTL and E-QTL detected by two-locus analysis was involved in Q × E and Q × Q × E interactions, supporting the results of ANOVA, where genotype × environment interaction were non-significant. The QTL for PHS detected in the present study may be efficiently utilized for marker-aided selection for enhancing PHST in bread wheat.     

Ethyloxanilates as specific male gametocides for wheat (Triticum aestivum L.)

Induction of male sterility by deployment of male gametocides holds immense potential in heterosis breeding of wheat. The efficacy of a new class of male gametocide for wheat (Triticum aestivum L.) is described: ethyloxanilates, the most active example of this class being ethyl 4‐fluorooxanilate (E4FO). E4FO induces male sterility, specifically, without detectable effects on various agronomic features and female fertility. The plants sprayed once with 0.15% E4FO exhibited 100% pollen and floret sterility without causing a significant reduction in total yield. E4FO was screened on 29 genotypes of wheat at 0.15% test concentration and was observed to induce 99.76 ± 0.37% male sterility. Thirteen F₁‐hybrids of wheat were produced using the gametocide in Winter 2000–2001 and were evaluated for their agronomic performance in Winter 2001–2002. The cross combinations viz., lines WR 544 × HW 2046 and HW 2044 × WR 956 have outperformed their respective better parents by 48.17% and 23.42% in grain yield/plant and thus have potential as hybrids.     

Molecular mapping of a dominant non-glaucousness gene from synthetic hexaploid wheat (Triticum aestivum L.)

The leaf and stem surfaces of many land plants are covered with a cuticular wax layer that confers a glaucous appearance or white bloom. Synthetic hexaploid wheat Line 3672 was non-glaucous, and common wheat Line 9753 was glaucous. The cuticular wax was characterized using Scanning Electron Microscopy (SEM). A hybrid using 9753 as female parent and 3672 as male parent was made and 108 F₂ plants and their F₃ progenies were used to map the non-glaucouness gene. Non-glaucousness in Line 3672 was controlled by a single dominant gene, temporarily designated Iw3672. Five SSR markers mapped on chromosome 2DS were linked to Iw3672. Additionally, two EST-derived markers and a SNP marker were developed and were also linked to Iw3672. The order of the eight markers and Iw3672 was Xte6 ₇₃₀ /Xbarc124 ₅₂₀ —Iw3672 —Xwe6 _2100/2150 —Xcau96 ₂₈₇ —Xcfd51 _180/200/230 —Xwe7—Xgdm5 ₁₉₀ —Xgdm35 _246/250 , with the genetic distance for each interval being 0.9 cM, 1.4 cM, 0.9 cM, 0.9 cM, 1.9 cM, 7.2 cM and 2.5 cM, respectively. We concluded that Iw3672 is physically mapped on the distal region of wheat chromosome 2DS.     

Genetic analysis of salt tolerance in a recombinant inbred population of wheat (Triticum aestivum L.)

A population of 114 recombinant inbred lines (RILs), derived from the cross Opata85 × W7984, was used to genetically analyze the response of wheat to salt stress. This analysis resulted in the identification of 47 QTL mapping to all wheat chromosomes except 1B, 1D, 4B, 5D and 7D. Of these QTL, 10 were effective during the germination stage, and 37 at the seedling stage. Many of the traits related to salt tolerance mapped to common chromosome intervals, such as Xglk683–Xcdo460 on chromosome 3A, Xfbb168–Xbcd147 on chromosome 3B, Xcdo1081–Xfbb226 on chromosome 4DL and Xpsr106–Xfbb283 on chromosome 6DL. QTL located in the interval Xcdo1081–Xfbb226 (chromosome 4DL) were effective during the germination stage, whereas those in the interval Xfbb231.1–Xmwg916 (chromosome 6DL) were relevant to the seedling stage. The QTL in the intervals Xglk683–Xcdo460 (chromosome 3AS) and Xfbb168–Xbcd147 (chromosome 3BL) were effective at both the germination and seedling stages.     

Genome-wide association study reveals loci associated with seed longevity in common wheat (Triticum aestivum L.)

Seed longevity could significantly determine seed regeneration cycle and greatly affect wheat production. With the 90 K chip assays, a genome-wide association study was performed to identify seed longevity-related markers and loci in common wheat. Seed germination ratios (GR) under artificially ageing of 166 wheat accessions across three environments were evaluated to assess seed longevity. Totally, 23 longevity-related loci were identified in the study, explaining 6.7%–11.4% of the phenotypic variations. Of these, QlgGR.cas-1A and QlgGR.cas-2B.2 were deemed as stable loci associated with wheat seed longevity. Fifteen loci were found overlapped with known quantitative trait loci or genes. Besides, QlgGR.cas-1A, QlgGR.cas-2B.2, QlgGR.cas-3D.1, QlgGR.cas-3D.2, QlgGR.cas-4A.2, QlgGR.cas-5A.1, QlgGR.cas-5A.2 and QlgGR.cas-6A.1 were colocated with seed dormancy-related loci. Significant additive effects were obtained for seed longevity by pyramiding favourable alleles. Several candidate genes were found involved in signal transduction and stress resistance pathways by sequencing analysis of significantly longevity-related molecular markers. These results might provide new sights into the genetic architecture of seed longevity.     

Genetic control of boron efficiency in wheat (Triticum aestivum L.)

The genetic control of boron (B) efficiencyin wheat (Triticum aestivum L.) wasstudied for three genotypes representing Binefficient (I, Bonza), moderately Binefficient (MI, SW 41) and B efficient (E,Fang 60) categories. Boron efficiency wasexpressed as a partially dominant characterbut the phenotypes of F₁ hybrids,relative to parents, indicated geneticcontrol varying from recessive to additiveto completely dominant with different crosscombinations and B levels. Major geneswere identified from the evaluation ofF₂-derived F₃ populations derivedfrom intercrosses between the threeparents. Monogenic segregation was foundin Bonza × SW 41 and SW 41 × Fang 60crosses and digenic segregation resultedin Bonza × Fang 60. Among thethree wheat genotypes with widely differentB efficiency, genetic variation forresponse to B could be accounted for by twogenes, Bo _d 1 and Bo _d 2.     

Limited but specific variations of seed storage proteins in Japanese common wheat (Triticum aestivum L.)

The electrophoretic banding patterns ofgliadin in common wheat lines derived fromJapan were determined byacid-polyacrylamide gel electrophoresis. For the 107 wheat lines used in our study,27 different patterns were identified, 13corresponding to the -gliadin, 8 tothe , -gliadin and 6 to the-gliadin. The gliadin patterns ofJapanese wheat cultivars and landracesgreatly differed from the patterns of wheatlines from other countries, and thevariation seen in wheat lines from Japanwas limited to 46 patterns. Sevencollection or breeding areas in Japanshowed different frequencies in theirgliadin patterns. Combining the gliadinpatterns with high molecular weightglutenin subunit compositions, 67combinations were observed. One gliadinpattern consisting of -gliadinpattern F, , -gliadinpattern H and -gliadin pattern Dwas frequently found in many Japanese wheatlines, though the other patterns werelimited to only one or two wheat lines.     

普通小麦氮素利用效率相关性状全基因组关联分析

氮元素在粮食作物生长和发育过程起着不可替代的作用。发掘氮素利用效率相关基因对于提升小麦产量、减少环境污染具有重要意义。植株根系构型(rootsystemarchitecture,RSA)代表着根系的结构及空间造型,显著受氮素水平影响。本研究在正常供氮和缺氮两种氮素水平下,对160份来自黄淮冬麦区和北部冬麦区普通小麦品系的根系构型相关性状(总根长、总根表面积、总根体积、平均根直径和根尖数)进行统计,并结合小麦660KSNP(single nucleotidepolymorphism)芯片基因分型数据对根系相关性状的相对值进行全基因组关联分析,以期发掘氮素利用效率相关位点。本研究检测到34个与氮素利用效率显著关联的SNP位点,可解释6.9%～15.4%的表型变异。关联位点在所有染色体均有分布,主要集中于1A、2B、3B、5B、6A、6B和7A染色体。11个位点与已报道位点重叠或接近,其他23个位点可能为新的位点。另外,在3B染色体上发现一个编码E3泛素连接酶的候选基因。     

普通小麦颖壳蜡质缺失突变体glossy1的转录组分析

为了明确突变体颖壳蜡质含量显著变化的分子机制,本研究对源自济麦22颖壳蜡质缺失突变体glossy1与野生型进行了转录组分析。结果表明,在glossy1突变体中,共筛选到12,230个差异表达基因,其中5811个基因在突变体中上调表达,6419个下调表达。GO(gene ontology)功能富集分析发现,差异基因主要富集在蜡质合成和转运途径,具体分布在酰基转移酶活性、脂质结合和水解酶活性等条目,由此推测这些途径与小麦穗部蜡质缺失性状是紧密相关的。我们还利用RT-qPCR检测了参与蜡质代谢途径部分基因的表达,结果与转录组结果是一致的。本研究为今后探究小麦蜡质代谢的分子机制和基因调控网络提供了数据支持,同时也为抗逆小麦育种奠定了理论基础。