Gene networks in hexaploid wheat: interacting quantitative trait loci for grain protein content

In hexaploid wheat, single-locus and two-locus quantitative trait loci (QTL) analyses for grain protein content (GPC) were conducted using two different mapping populations (PI and PII). Main effect QTLs (M-QTLs), epistatic QTLs (E-QTLs) and QTL x environment interactions (QE, QQE) were detected using two-locus analyses in both the populations. Only a few QTLs were common in both the analyses, and the QTLs and the interactions detected in the two populations differed, suggesting the superiority of two-locus analysis and the need for using several mapping populations for QTL analysis. A sizable proportion of genetic variation for GPC was due to interactions (28.59% and 54.03%), rather than to M-QTL effects (7.24% and 7.22%), which are the only genetic effects often detected in the majority of QTL studies. Even E-QTLs made a marginal contribution to genetic variation (2.68% and 6.04%), thus suggesting that the major part of genetic variation is due to changes in gene networks rather than the presence or absence of specific genes. This is in sharp contrast to the genetic dissection of pre-harvest sprouting tolerance conducted by us earlier, where interaction effects were not substantial, suggesting that the nature of genetic variation also depends on the nature of the trait.     

Identification for quantitative trait loci controlling grain shattering in rice

Seed shattering is an important factor causing loss of grain yield before and during rice harvest. In the present study, the quantitative trait loci regarding shattering scale, breaking tensile strength (BTS) and abscission layer (AL), the parameters evaluating seed shattering habit by hand gripping, a digital force gauge and observation on AL, respectively, were identified by using an doubled haploid line (DHL) population from a cross between a loose-shattering type Tongil variety, ‘Samgang’, and a moderately difficult shattering japonica variety, ‘Nagdong’. Eight QTLs consisted in four QTLs for shattering scale, two QTLs for AL, each one QTL for pulling and bending strength were detected on six chromosomes, respectively. Among them, Qss1 with flanking markers RM6696 and RM476 explained 31% of phenotype variation in shattering scale. Furthermore, two new QTLs controlling shattering habit, Qss5-2 and Qal5-1, were located on chromosome 5 at the interval 5028–5037 and 5021-RM289. They explained 10% and 12% of phenotype variations, respectively. A total of eleven digenic epistatic loci were identified for four parameters. The identification of QTLs affecting seed shattering habits is favorable to thoroughly dissect the genetic mechanism of the shattering habit and to apply for marker-assisted selection in rice breeding system of specific regions.     

Identification of quantitative trait loci for cadmium tolerance and accumulation in wheat

Quantitative trait loci (QTL) for Cadmium (Cd) tolerance and accumulation in wheat (Triticum aestivum L.) were identified, using 103 recombinant inbred lines (RILs) derived from a cross of Ch × Sh at germination and seedling stages. The traits of germination, growth and physiology were measured. Cd tolerance indexes (TI) were calculated for plants under Cd stress relative to control conditions. Cd concentrations in both root and shoot were determined and the amount of Cd accumulation and translocation calculated. The phenotypic variation of the above traits showed a continuous distribution pattern among the RILs. Twenty-six QTLs were detected, (16 of which were designated for the traits under the control and Cd stress, 8 for Cd tolerance and 2 for root Cd accumulation). These 26 QTLs individually could explain 7.97–60.16% of the phenotypic variation. Fourteen QTLs were positive (with the additive effects coming from Ch) while the remaining 12 QTLs were negative (with the additive effects contributed by Sh). No QTL were detected in the same region on the chromosomes of wheat. The results indicated that genetic mechanisms controlling the traits of Cd tolerance were independent from each other. Therefore, in this study, the properties of Cd tolerance and accumulation showed to be independent traits in wheat.     

Identification of quantitative trait loci for production traits in commercial pig populations

The aim of this study was to investigate methods for detecting QTL in outbred commercial pig populations. Several QTL for back fat and growth rate, previously detected in experimental resource populations, were examined for segregation in 10 different populations. Two hundred trait-by-population-by-chromosome tests were performed, resulting in 20 tests being significant at the 5% level. In addition, 53 QTL tests for 11 meat quality traits were declared significant, using a subset of the populations. These results show that a considerable amount of phenotypic variance observed in these populations can be explained by major alleles segregating at several of the loci described. Thus, despite a relatively strong selection pressure for growth and back fat traits in these populations, these alleles have not yet reached fixation. The approaches used here demonstrate that it is possible to verify segregation of QTL in commercial populations by limited genotyping of a selection of informative animals. Such verified QTL may be directly exploited in marker-assisted selection (MAS) programs in commercial populations and their molecular basis may be revealed by positional candidate cloning.     

Identification and validation of quantitative trait loci conferring tan spot resistance in the bread wheat variety Ernie

Tan spot, caused by Pyrenophora tritici-repentis, is a foliar disease of wheat, and it can inflict serious reduction in grain yield and quality. The bread wheat variety Ernie was found to be immune to this disease in Australia, and its genetic control was investigated by quantitative trait loci (QTL) analysis using a doubled haploid population. Eight QTL were identified in this population from three independent trials, and four of them were derived from the parent Ernie. The most significant QTL was located on chromosome arm 2BS, explaining 38.2, 29.8 and 36.2% of the phenotypic variance, respectively, in these trials. The effects of the 2BS QTL were further validated in four additional populations. The presence of this single QTL reduced disease severity by between 29.2 and 67.1% with an average of 50.5%. The significant effects of this QTL and its consistent detection across all the trials with different genetic backgrounds make it an ideal target for breeding programmes as well as for its further characterization. Data from this study also showed that neither plant height nor heading date significantly affects tan spot resistance.     

Identification and mapping of leaf, stem and stripe rust resistance quantitative trait loci and their interactions in durum wheat

Leaf rust (Puccinia triticina Eriks.), stripe rust (Puccinia striiformis f. tritici Eriks.) and stem rust (Puccinia graminis f. sp. tritici) cause major production losses in durum wheat (Triticum turgidum L. var. durum). The objective of this research was to identify and map leaf, stripe and stem rust resistance loci from the French cultivar Sachem and Canadian cultivar Strongfield. A doubled haploid population from Sachem/Strongfield and parents were phenotyped for seedling reaction to leaf rust races BBG/BN and BBG/BP and adult plant response was determined in three field rust nurseries near El Batan, Obregon and Toluca, Mexico. Stripe rust response was recorded in 2009 and 2011 nurseries near Toluca and near Njoro, Kenya in 2010. Response to stem rust was recorded in field nurseries near Njoro, Kenya, in 2010 and 2011. Sachem was resistant to leaf, stripe and stem rust. A major leaf rust quantitative trait locus (QTL) was identified on chromosome 7B at Xgwm146 in Sachem. In the same region on 7B, a stripe rust QTL was identified in Strongfield. Leaf and stripe rust QTL around DArT marker wPt3451 were identified on chromosome 1B. On chromosome 2B, a significant leaf rust QTL was detected conferred by Strongfield, and at the same QTL, a Yr gene derived from Sachem conferred resistance. Significant stem rust resistance QTL were detected on chromosome 4B. Consistent interactions among loci for resistance to each rust type across nurseries were detected, especially for leaf rust QTL on 7B. Sachem and Strongfield offer useful sources of rust resistance genes for durum rust breeding.     

Identification of quantitative trait loci for physical and chemical properties of rice grain

Quantitative trait loci (QTL) associated with six physical traits of cooked rice and seven chemical properties of rice grain were identified using a recombinant inbred (RI) population of rice evaluated over 3 years at the National Honam Agricultural Research Institute in Korea. The RI population consisted of 164 lines derived from a cross between Milyang23 and Gihobyeo, and the genetic map consisted of 414 molecular markers. A total of 49 QTL were identified for the 13 physico-chemical properties using composite interval mapping. Of these, 13 QTL were identified for 2 or more years, while 36 were detected in only 1 year. Five QTL were identified over all 3 years and will be useful for marker-assisted improvement of rice grain quality in Korea. The two QTL with the highest LOD scores, adhesiveness1.2 and potassium content7.1, provide a valuable starting point for positional cloning of genes underlying these QTL.     

Identification and validation of quantitative trait loci for seed yield,oil and protein contents in two recombinant inbred line populations of soybean

Soybean seeds contain high levels of oil and protein, and are the important sources of vegetable oil and plant protein for human consumption and livestock feed. Increased seed yield, oil and protein contents are the main objectives of soybean breeding. The objectives of this study were to identify and validate quantitative trait loci (QTLs) associated with seed yield, oil and protein contents in two recombinant inbred line populations, and to evaluate the consistency of QTLs across different environments, studies and genetic backgrounds. Both the mapping population (SD02-4-59 × A02-381100) and validation population (SD02-911 × SD00-1501) were phenotyped for the three traits in multiple environments. Genetic analysis indicated that oil and protein contents showed high heritabilities while yield exhibited a lower heritability in both populations. Based on a linkage map constructed previously with the mapping population and using composite interval mapping and/or interval mapping analysis, 12 QTLs for seed yield, 16 QTLs for oil content and 11 QTLs for protein content were consistently detected in multiple environments and/or the average data over all environments. Of the QTLs detected in the mapping population, five QTLs for seed yield, eight QTLs for oil content and five QTLs for protein content were confirmed in the validation population by single marker analysis in at least one environment and the average data and by ANOVA over all environments. Eight of these validated QTLs were newly identified. Compared with the other studies, seven QTLs for seed yield, eight QTLs for oil content and nine QTLs for protein content further verified the previously reported QTLs. These QTLs will be useful for breeding higher yield and better quality cultivars, and help effectively and efficiently improve yield potential and nutritional quality in soybean.     

Identification of quantitative trait loci for race-nonspecific resistance to tan spot in wheat

Tan spot, caused by Pyrenophora tritici-repentis (Ptr), is an economically important foliar disease in the major wheat growing areas throughout the world. Multiple races of the pathogen have been characterized based on their ability to cause necrosis and/or chlorosis on differential wheat lines. In this research, we evaluated a population of recombinant inbred lines derived from a cross between the common wheat varieties Grandin and BR34 for reaction to tan spot caused by Ptr races 1–3 and 5. Composite interval mapping revealed QTLs on the short arm of chromosome 1B and the long arm of chromosome 3B that were significantly associated with resistance to all four races. The effects of the two QTLs varied for the different races. The 1B QTL explained from 13% to 29% of the phenotypic variation, whereas the 3B QTL explained from 13% to 41% of the variation. Additional minor QTLs were detected but not associated with resistance to all races. The host-selective toxin Ptr ToxA, which is produced by races 1 and 2, was not a significant factor in the development of disease in this population. The race-nonspecific resistance derived from BR34 may take precedence over the gene-for-gene interaction known to be associated with the wheat–Ptr system.     

Mapping quantitative trait loci in tetraploid populations

Knowledge of quantitative trait locus (QTL) mapping in polyploids is almost void, albeit many exquisite strategies of QTL mapping have been proposed and extensive investigations have been carried out in diploid animals and plants. In this paper we develop a simple algorithm which uses an iteratively reweighted least square method to map QTLs in tetraploid populations. The method uses information from all markers in a linkage group to infer the probability distribution of QTL genotype under the assumption of random chromosome segregation. Unlike QTL mapping in diploid species, here we estimate and test the compound 'gametic effect', which consists of the composite 'genic effect' of alleles and higher-order gene interactions. The validity and efficiency of the proposed method are investigated through simulation studies. Results show that the method can successfully locate QTLs and separates different sources (e.g. additive and dominance) of variance components contributed by the QTLs.     

Development and validation of molecular markers for grain cadmium in durum wheat

Durum wheat is capable of accumulating cadmium, a toxic heavy metal, in the grain at levels that have been deemed unsafe for human consumption. Previous studies have identified genetic variation as well as markers associated with Cd accumulation in durum wheat, which can be exploited to develop low Cd cultivars. Because the phenotyping for Cd content is very expensive, KASP markers were developed from molecular markers associated with grain Cd and tested for their usefulness for marker-assisted breeding. A total of 1278 unique genotypes from preliminary and advanced yield trials grown at multiple locations for 2 years were evaluated for grain Cd as well as screened for markers associated with Cd uptake. One marker on chromosome 5B was polymorphic in all crosses between high and low Cd parents and had r² values ranging from 0.38 to 0.85. Two other markers on the same chromosome predicted similar levels of variation in many trials; however, they were not polymorphic in all populations. The KASP markers accurately predicted up to 97% of the lines for Cd phenotype in different trials. This study identified two markers, Cad-5B and Ex_c1343_2570756, with an average prediction accuracy of 84–88%. These markers could be useful for marker-assisted selection for low grain Cd in durum wheat.     

Quantitative trait loci for grain yield and adaptation of durum wheat (Triticum durum Desf.) across a wide range of water availability

Grain yield is a major goal for the improvement of durum wheat, particularly in drought-prone areas. In this study, the genetic basis of grain yield (GY), heading date (HD), and plant height (PH) was investigated in a durum wheat population of 249 recombinant inbred lines evaluated in 16 environments (10 rainfed and 6 irrigated) characterized by a broad range of water availability and GY (from 5.6 to 58.8 q ha(-1)). Among the 16 quantitative trait loci (QTL) that affected GY, two major QTL on chromosomes 2BL and 3BS showed significant effects in 8 and 7 environments, with R2 values of 21.5 and 13.8% (mean data of all 16 environments), respectively. In both cases, extensive overlap was observed between the LOD profiles of GY and PH, but not with those for HD. QTL specific for PH were identified on chromosomes 1BS, 3AL, and 7AS. Additionally, three major QTL for HD on chromosomes 2AS, 2BL, and 7BS showed limited or no effects on GY. For both PH and GY, notable epistasis between the chromosome 2BL and 3BS QTL was detected across several environments.     

Mapping and validation of quantitative trait loci associated with wheat yellow mosaic bymovirus resistance in bread wheat

Wheat yellow mosaic (WYM) caused by wheat yellow mosaic bymovirus (WYMV) has been growing as one of the most serious diseases affecting wheat production in China. In this study, the association of quantitative trait loci (QTLs) governing WYMV resistance with molecular markers was established using 164 recombinant inbred lines (RILs) derived from 'Xifeng Wheat' (highly resistant)?×?'Zhen 9523' (highly susceptible). Phenotypic data of WYMV resistance of the RILs were collected from 4-year, two-location replicated field trials. A molecular marker-based linkage map, which was comprised of 273 non-redundant loci and represented all the 21 wheat chromosomes, was constructed with the JoinMap 4.0 software. Using the Windows QTL Cartographer V2.5 software, three QTLs associated with WYMV resistance, QYm.njau-3B.1, QYm.njau-5A.1 and QYm.njau-7B.1, were detected on chromosomes 3BS, 5AL, and 7BS, respectively. The favorable allele effects were all contributed by 'Xifeng Wheat'. Among the three QTLs, QYm.njau-3B.1 and QYm.njau-5A.1 were detected in all the four trials and the overall mean, and could explain 3.3-10.2% and 25.9-53.7% of the phenotypic variation, respectively, while QYm.njau-7B.1 was detected in one trial and the overall mean and explained 4.9 and 3.3% of the phenotypic variation, respectively. A large portion of the variability for WYMV response was explained by a major QTL, QYm.njau-5A.1. The relationship of the molecular markers linked with QYm.njau-5A.1 and the WYMV resistance was further validated using a secondary F(2) population. The results showed that three markers, i.e., Xwmc415.1, CINAU152, and CINAU153, were closely linked to QYm.njau-5A.1 with the genetic distances of 0.0, 0.0, and 0.1?cM, respectively, indicating they should be useful in marker-assisted selection (MAS) wheat breeding for WYMV resistance. A panel of germplasm collection consisting of 46 wheat varieties with known WYMV response phenotypes was further used to validate the presence and effects of QYm.njau-5A.1 and the above three markers. It was found that QYm.njau-5A.1 was present in 12 of the 34 WYMV-resistant varieties.     

Detection of QTLs for grain protein content in durum wheat

Grain protein content (GPC) of durum wheat (Triticum turgidum L. var. durum) is an important trait for the nutritional value of grain and for influencing the technological property of flour. Protein content is a quantitative trait negatively correlated with grain yield, thus increase in protein quantity usually results in yield reduction. This study was initiated to introgress alleles for high GPC from var. dicoccoides into durum wheat germplasm by the backcross inbred line (BIL) method and to identify molecular markers linked to high GPC alleles not associated with depressing effects on yield. The backcross line 3BIL-85 with high GPC and similar grain yield to the recurrent parent was backcrossed to Latino, and the generations F₂, F₃ and F₄ were evaluated for GPC and yield per spike (GYS) in three field trials. Three QTLs with major effects on GPC were detected on chromosome arms 2AS, 6AS and 7BL, identified by the markers Xcfa2164, XP39M37 ₍₂₅₀₎ and Xgwm577 _, respectively. Multiple regression analysis indicated that the three QTLs explained all the genetic variances of the trait. The high GPC parental line 3BIL-85 was not significantly different from the recurrent parent Latino for GYS, but the phenotypic correlation coefficient between GPC and GYS had negative values (from −0.02 to −0.28) in each trial, although it was statistically significant only in the F3 progeny trial. No co-located QTL for GYS was detected, excluding the hypothesis that the putative QTLs for GPC were indirect QTLs for low grain yield. The negative protein-yield response could be due to: (a) co-location of grain yield per spike QTLs with reduced phenotypic effects not detectable by the experimental design or statistical procedures, or to (b) opposite pleiotropic gene effects due to the major bio-energetic requirements for synthesis of protein then carbohydrates. Mapping loci by BILs should enable the production of near-isogenic lines in which the individual effects of each QTL can be examined in detail without confounding variations due to other putative QTLs. An erratum to this article can be found at     

Identification of quantitative trait loci and associated candidate genes for low-temperature tolerance in cold-hardy winter wheat

Low-temperature (LT) tolerance is an important economic trait in winter wheat (Triticum aestivum L.) that determines the plants’ ability to cope with below freezing temperatures. Essential elements of the LT tolerance mechanism are associated with the winter growth habit controlled by the vernalization loci (Vrn-1) on the group 5 chromosomes. To identify genomic regions, which in addition to vrn-1 determine the level of LT tolerance in hexaploid wheat, two doubled haploid (DH) mapping populations were produced using parents with winter growth habit (vrn-A1, vrn-B1, and vrn-D1) but showing different LT tolerance levels. A total of 107 DH lines were analyzed by genetic mapping to produce a consensus map of 2,873 cM. The LT tolerance levels for the Norstar (LT₅₀=−20.7°C) × Winter Manitou (LT₅₀=−14.3°C) mapping population ranged from −12.0 to −22.0°C. Single marker analysis and interval mapping of phenotyped lines revealed a major quantitative trait locus (QTL) on chromosome 5A and a weaker QTL on chromosome 1D. The 5A QTL located 46 cM proximal to the vrn-A1 locus explained 40% of the LT tolerance variance. Two C-repeat Binding Factor (CBF) genes expressed during cold acclimation in Norstar were located at the peak of the 5A QTL.