Advanced-backcross QTL analysis in spring barley: IV. Localization of QTL × nitrogen interaction effects for yield-related traits

The advanced backcross quantitative trait locus (AB-QTL) analysis has proven its usefulness to identify and localize favourable alleles from exotic germplasm and to transfer those alleles into elite varieties. In a balanced design with up to six environments and two nitrogen fertilization (N treatment) levels, a 4-factorial mixed model analysis of variance (ANOVA) was used to identify QTL main effects, QTL × environment interaction effects and QTL × N treatment interaction effects in the spring barley BC₂DH population S42. The yield-related traits studied were number of ears per m², days until heading, plant height, thousand grain weight (TGW) and grain yield. In total, 82 QTLs were detected for all traits. This finding was compared to a previous QTL study of the same population S42, where the current field data was reduced to one half through restriction of the analysis to the standard N treatment level (von Korff et al., Theor Appl Genet 112: 1221–1231, 2006). These authors located 54 QTLs for the same traits by applying a 3-factorial mixed model similar to the current model but excluding the factor N treatment. We found that QTL × environment interaction, alone or in combination, accounted for 24 of the newly uncovered QTLs, whereas QTL × N treatment interaction was of lesser importance with six new cases in total. A valuable QTL interacting with N treatment has been identified on chromosome 7H where lines carrying the wild barley allele were superior in number of ears per m² in either N treatment. We conclude that in population S42 the extension of the phenotype data set and the inclusion of N treatment into the mixed model increased the power of QTL detection by providing an additional replication rather than by revealing specific N treatment QTLs.     

Correction: QTL mapping under salt stress in rice using a Kalarata–Azucena population

Euphytica - The color of flour and its end-use products is an important quality trait of wheat. Understanding the genetic basis of this trait is essential for improving wheat quality. In this...     

Comparison and analysis of main effects,epistatic effects,and QTL × environment interactions of QTLs for agronomic traits using DH and RILs populations in rice

Two genetic linkage maps based on doubled haploid (DH) and recombinant inbred lines (RILs) populations, derived from the same indica-japonica cross ‘Samgang × Nagdong’, were constructed to analyze the quantitative trait loci (QTLs) affecting agronomic traits in rice. The segregations of agronomic traits in RILs population showed larger variations than those in DH population. A total of 10 and 12 QTLs were identified on six chromosomes using DH population and seven chromosomes using RILs population, respectively. Three stable QTLs including pl9.1, ph1.1, and gwp11.1 were detected through different years. The percentages of phenotypic variation explained by individual QTLs ranged from 8 to 18% in the DH population and 9 to 33% in the RILs population. Twenty-three epistatic QTLs were identified in the DH population, while 21 epistatic QTLs were detected in the RILs population. Epistatic interactions played an important role in controlling the agronomic traits genetically. Four significant main-effect QTLs were involved in the digenic interactions. Significant interactions between QTLs and environments (QE) were identified in two populations. The QTLs affecting grain weight per panicle (GWP) were more sensitive to the environmental changes. The comparison and QTLs analysis between two populations across different years should help rice breeders to comprehend the genetic mechanisms of quantitative traits and improve breeding programs in marker-assisted selection (MAS).     

Genetic variation,genotype × environment interaction,and selection for tipburn resistance in lettuce in multi-environments

Tipburn is a calcium related and environmentally induced physiological disorder causing economic damage in all lettuce (Lactuca sativa L.) production regions. The objectives of this research were to determine (1) the genetic variation for tipburn incidence, (2) the genotype (G) × environment (E) interaction (GE) for tipburn incidence, and (3) the efficiency of field selection for tipburn resistance. Tipburn incidence was recorded over 2 years in Salinas, CA, and Yuma, AZ, for 55 romaine, crisphead, green leaf, and red leaf type cultivars, and over 3 years in Quebec for 15 romaine cultivars. Analysis revealed that G, E, and GE affected tipburn incidence, including crossover interactions that were not repeatable over years. This indicates that cultivar/breeding line evaluations should be based on mean performance and stability over multiple environments. Among lettuce types, only crisphead had significant genetic variability for tipburn resistance, reflecting the greater breeding effort applied to this type compared romaine, green and red leaf types. Analysis of a dataset with five romaine cultivars in eight environments in California, Arizona, and Quebec for 2 years revealed that Yuma in 2006 and Saint-Blaise in 2005 were highly correlated (r = 0.923, P < 0.05), and were the most discriminating and most representative environments for tipburn evaluation. Single plant selection for tipburn resistance in three F₂ romaine populations was ineffective. Further, the degree of head closure was significantly associated with tipburn incidence. Identification and selection of morphological characters associated with resistance in conjunction with direct selection against tipburn may be an effective method for genetic improvement of tipburn resistance.     

Genetic effects and genotype × environment interactions for cooking quality traits in Indica-japonica crosses of rice (Oryza sativa L.)

Indica-japonica hybridization is an important approach for developing superior performing hybrids in rice (Oryza sativa L.). In view of the scanty information available on cooking quality characters in indica-japonica crosses, an investigation was undertaken to estimate genetic and genotype × environment variance and covariance components of amylose content, gel consistency and alkali digestion value, and to determine the relative importance of direct genetic effects, maternal genetic effects and cytoplasmic effects in the genetic variations of the three quality characters. Two indica photo-sensitive genic male sterile (PGMS) lines and four japonica varieties were used as parents to make crosses. Genetic model with genotype × environment interactions for triploid endosperm was used for genetic studies of the three cooking quality characters. Variance component analysis revealed that genetic variations of the three characters were mainly attributable to direct additive and maternal additive effects, and the three traits had significant direct and maternal heritabilities. Genotype × environment interactions were mainly dominance × environment (including direct dominance × environment and maternal dominance × environment) and cytoplasm × environment interactions. Environment factors could only affect the expression extent of dominant genes, without changing their directions. Predicted values of genetic effects indicated that the parental lines, ‘VI-70’ and ‘H9304-1’, appeared to be best for amylose content, ‘T 1950’ and ‘Suxuan’ appeared to be best for gel consistency and alkali digestion value. This revised version was published online in August 2006 with corrections to the Cover Date.     

Mapping quantitative trait loci (QTL) for grain size in rice using a RIL population from Basmati × indica cross showing high segregation distortion

Grain size is one of the three productivity related traits in rice and hence a major target for genetic improvement. Since understanding genetic variation in grain size between Basmati and indica genotypes is important for rice improvement, a recombinant inbred population was developed from a traditional aromatic cultivar ‘Basmati 370’ and a non-aromatic indica genotype ‘IRBB60’. This population was phenotyped in two locations for grain length (GL), grain breadth (GB), GL/GB ratio (LBR) and grain weight (GW). Though the RIL population reported in the current study exhibited segregation distortion (SD) for 54 % of the markers, they were utilized in analysis using an appropriate statistical package, PROC QTL in the SAS environment. Interval mapping revealed a total of 15 QTLs for GL, seven for GB, 15 for LBR and two for GW. Among them 13 were not reported earlier and thus novel. For a known major QTL identified in the study, GW8 for GB, a PCR based functional marker was designed and validated. This is the first report wherein a very high proportion of markers (>50 %) exhibiting SD have been successfully used for QTL mapping.     

QTL analysis for eating quality-related traits in an F2:3 population derived from waxy corn × sweet corn cross

In order to identify quantitative trait loci (QTL) for the eating quality of waxy corn and sweet corn (Zea mays L.), QTL analysis was conducted on an F₂ population derived from a cross between a waxy corn inbred line and a sweet corn inbred line. Ten QTLs for pericarp thickness (PER), amylose content (AMY), dextrose content (DEX) and sucrose content (SUC) were found in the 158 F₂ families. Among them, four QTLs, qAMY4 (10.43%), qAMY9 (19.33%), qDEX4 (21.31%) and qSUC4 (30.71%), may be considered as major QTLs. Three of these, qAMY4, qDEX4 and qSUC4, were found to be located within a region flanked by two adjacent SSR markers on chromosome 4 (umc1088 and bnlg1265), making this SSR marker pair a useful selection tool for screening the eating quality traits of AMY, DEX and SUC. The QTL for amylose content was found to be located between markers phi027 and umc1634, raising the possibility of its identity being the Wx1 gene, which encodes a granule-bound amylose synthase. The new QTLs identified by the present study could serve as useful molecular markers for selecting important eating quality traits in subsequent waxy corn breeding studies.     

Inheritance of anthocyanin content in the ripe berries of a tetraploid × diploid grape cross population

Variation patterns and inheritance of anthocyanin content in the ripe berries of a tetraploid × diploid table grape cross population were investigated in two successive years. The population segregated for three different ploid levels: dipolids, triploids, and tetraploids. A total of 28 different anthocyanins were detected and quantified in the progeny population. Transgressive segregation for the total anthocyanin content was observed in all the three ploid progeny populations. The total anthocyanin content increased as the ploid level increased. The broad sense heritabilities (H²) of the total anthocyanin content were all relatively high, ranging from 0.53 to 0.98, 0.57 to 0.97 and 0.43 to 0.94 in the diploid, triploid and tetraploid population, respectively. Our results suggested that the total anthocyanin content followed an additive inheritance model in this polyploid segregation population. We also observed that the relative contribution of individual anthocyanins to the total anthocyanin content varied significantly among different ploid populations, suggesting that genetic background has important impact on the accumulation of the individual anthocyanin compounds. These results will help develop better breeding strategies in a polyploid table grape breeding program for improving the content of anthocyanins, an important class of polyphenolics possessing antioxidant activities and many other health-related benefits.     

Quantitative trait locus mapping for Verticillium wilt resistance in a backcross inbred line population of cotton (Gossypium hirsutum × Gossypium barbadense) based on RGA-AFLP analysis

Verticillium wilt (VW), caused by Verticillium dahliae Kleb., is one of the most important diseases in cotton. The objective of this study was to map quantitative trait loci (QTLs) conferring VW resistance using resistance gene analog (RGA)-targeted amplified fragment length polymorphism (RGA-AFLP) markers in an interspecific backcross inbred line mapping population, consisting of 146 lines from a susceptible Sure-Grow 747 (Gossypium hirsutum L.) × resistant Pima S-7 (G. barbadense L.) cross. VW resistance was evaluated in replicated tests based on disease incidence in the field, and disease incidence and severity in the greenhouse. Of 160 polymorphic RGA-AFLP markers, 42 were significantly correlated with one or more VW traits and 41 were placed on a linkage map which covered 1,226 cM of the cotton genome and contained 251 other molecular markers. Three QTLs for VW resistance were detected, each of which explained 12.0–18.6 % of the phenotypic variation. Two of these QTLs for disease incidence and severity detected in the greenhouse inoculation tests using root wounding are located on chromosome c4. Both are closely linked to four RGA-AFLP markers and therefore considered as the same QTL for VW resistance. The other QTL detected in the field test was located on c19 and flanked by several RGA-AFLP markers. The desirable QTL allele on c4 for VW resistance detected in the greenhouse was from the VW susceptible Upland parent and absent from the resistant Pima parent which was more VW susceptible due to the disarmament of the first line of defense mechanism due to root wounding during inoculation. The other desirable VW resistance QTL allele, on c19, was from the resistant parent Pima S-7, consistent with the fact that Pima cotton was more resistant to VW when naturally infected in the field. The results should facilitate the development of more sequence specific markers and the transfer of VW resistance from Pima to Upland cotton through marker-assisted selection.     

Genotype × environment interaction and genetic improvement for yield and yield stability of rainfed durum wheat in Iran

The genotype × environment (GE) interaction influences genotype selection and recommendations. Consequently, the objectives of genetic improvement should include obtaining genotypes with high potential yield and stability in unpredictable conditions. The GE interaction and genetic improvement for grain yield and yield stability was studied for 11 durum breeding lines, selected from Iran/ICARDA joint program, and compared to current checks (i.e., one durum modern cultivar and two durum and bread wheat landraces). The genotypes were grown in three rainfed research stations, representative of major rainfed durum wheat-growing areas, during 2005–09 cropping seasons in Iran. The additive main effect and multiplicative interaction (AMMI) analysis, genotype plus GE (GGE) biplot analysis, joint regression analysis (JRA) (b and S²di), six stability parameters derived from AMMI model, two Kang’s parameters [i.e., yield-stability (YSi) statistic and rank-sum], GGE distance (mean performance + stability evaluation), and two adaptability parameters [i.e., TOP (proportion of environments in which a genotype ranked in the top third) and percentage of adaptability (Ad)] were used to analyze GE interaction in rainfed durum multi-environment trials data. The main objectives were to (i) evaluate changes in adaptation and yield stability of the durum breeding lines compared to modern cultivar and landraces (ii) document genetic improvement in grain yield and analyze associated changes in yield stability of breeding lines compared to checks and (iii) to analyze rank correlation among GGE biplot, AMMI analysis and JRA in ranking of genotypes for yield, stability and yield-stability. The results showed that the effects due to environments, genotypes and GE interaction were significant (P < 0.01), suggesting differential responses of the genotypes and the need for stability analysis. The overall yield was 2,270 kg ha^?1 for breeding lines and modern cultivar versus 2,041 kg ha^?1 for landraces representing 11.2 % increase in yield. Positive genetic gains for grain yield in warm and moderate locations compared to cold location suggests continuing the evaluation of the breeding material in warm and moderate conditions. According to Spearman’s rank correlation analysis, two types of associations were found between the stability parameters: the first type included the AMMI stability parameters and joint regression parameters which were related to static stability and ranked the genotypes in similar fashion, whereas the second type consisted of the rank-sum, YSi, TOP, Ad and GGED which are related to dynamic concept of stability. Rank correlations among statistical methods for: (i) stability ranged between 0.27 and 0.97 (P < 0.01), was the least between AMMI and GGE biplot, and highest for AMMI and JRA and (ii) yield-stability varied from 0.22 (between GGE and JRA) to 0.44 (between JRA and AMMI). Breeding lines G8 (Stj3//Bcr/Lks4), G10 (Ossl-1/Stj-5) and G12 (modern cultivar) were the best genotypes in terms of both nominal yield and stability, indicating that selecting for improved yield potential may increase yield in a wide range of environments. The increase in adaptation, yield potential and stability of breeding lines has been reached due to gradual accumulation of favorable genes through targeted crosses, robust shuttle breeding and multi-locational testing.     

Changes in the concept of genotype × environment interactions to fit agriculture diversification and decentralized participatory plant breeding: pluridisciplinary point of view

The standardization of environments (E) encouraged by modern society and by the productivist model of agriculture has resulted in the standardization of genotypes (G) thereby reducing G × E interaction. New societal values call for the diversification of agriculture to fit contrasted environments. This process can be depicted by four models defined by two axes, one socio-economic (individual logics versus collective governance), and the other agro-ecological (reductionist versus systemic approaches). These models differ in (i) their objectives (from improvement in yield to the empowerment of farmers), (ii) their specific expectations with respect to genotypes (from inherited genetic resources to varieties that represent genetic, ethical and social progress), (iii) their specific representations of the environment (E) (from a simple interaction between the bio-physical environment (B) and the crop management (C), to a complex interaction including the competences of the actors (A), outlets (O), regulations (R), society (S)), (iv) their particular relations between G and E (from G × E to G × B × C × A under evolving constraints represented by R × O × S). Taking this diversity into account changes the way plant improvement is considered. Thus, depending on the model, the order, interest and status of the five classic stages of plant improvement (setting objectives, creating variability, selecting, evaluating and disseminating) may be called into question. Between the existing analytical model (Model I) and a holistic model (Model IV) which remains to be developed, lies the challenge of ensuring the sustainability, efficiency and acceptability of plant breeding and resulting innovations. From a simple “statistical parameter” that we, as plant breeders, attempt to reduce, the G × E interaction is becoming an “objective” that we try to predict and valorize. Structuring the different components of E, G and G × E, enables us to extend the basic concept of representivity to both the cultivation conditions and the relational socio-economic positions of the actors involved.

Advanced backcross QTL analysis reveals complicated genetic control of rice grain shape in a japonica × indica cross

Grain shape is an important trait for improving rice yield. A number of quantitative trait loci (QTLs) for this trait have been identified by using primary F₂ mapping populations and recombinant inbred lines, in which QTLs with a small effect are harder to detect than they would be in advanced generations. In this study, we developed two advanced mapping populations (chromosome segment substitution lines [CSSLs] and BC₄F₂ lines consisting of more than 2000 individuals) in the genetic backgrounds of two improved cultivars: a japonica cultivar (Koshihikari) with short, round grains, and an indica cultivar (IR64) with long, slender grains. We compared the ability of these materials to reveal QTLs for grain shape with that of an F₂ population. Only 8 QTLs for grain length or grain width were detected in the F₂ population, versus 47 in the CSSL population and 65 in the BC₄F₂ population. These results strongly suggest that advanced mapping populations can reveal QTLs for agronomic traits under complicated genetic control, and that DNA markers linked with the QTLs are useful for choosing superior allelic combinations to enhance grain shape in the Koshihikari and IR64 genetic backgrounds.     

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.     

Molecular mapping of quantitative trait loci for domestication traits and β-glucan content in a wheat recombinant inbred line population

Genetic maps are useful for analysis of quantitative trait loci (QTLs) and for marker-assisted selection (MAS) in breeding. A simple sequence repeat (SSR) marker linkage map of common wheat was constructed based on recombination inbred lines (RILs) derived from a cross between Chinese Spring and spelt wheat. The map included 264 loci on all wheat chromosomes covering 2,345.2 cM with 962, 794.6, and 588.6 cM for the A, B, and D genomes, respectively. Using the RILs and the map, we detected 42 putative QTLs on 15 chromosomes for ear length, spikelet number, spike compactness, kernel length, kernel width, kernel height and β-glucan content. Each QTL explained 4–45% of the phenotypic variation. Five QTL cluster regions were detected on chromosomes 1A, 5AL, 2B, 2D, and 4D. The first QTLs for β-glucan content in wheat were identified on chromosomes 3A, 1B, 5B, and 6D.     

Inheritance of sugar and acid contents in the ripe berries of a tetraploid × diploid grape cross population

Pearl millet [Pennisetum glaucum (L.) R. Br.] is a major warm-season cereal, grown primarily for grain production in the arid and semi-arid tropical regions of Asia and Africa. Iron (Fe) and zinc (Zn) deficiencies have been reported to be a food-related primary health problem affecting nearly two billion people worldwide. Improving Fe and Zn densities of staple crops by breeding offers a cost-effective and sustainable solution to reducing micronutrient malnutrition in resource poor communities. An understanding of the genetics of these micronutrients can help to accelerate the breeding process, but little is known about the genetics and heterosis pattern of Fe and Zn densities in pearl millet. In the present study, ten inbred lines and their full diallel crosses were used to study the nature of gene action and heterosis for these micronutrients. The general combining ability (GCA) effects of parents and specific combining ability (SCA) effects of hybrids showed significant differences for both of the micronutrients. However, the predictability ratio (2σ²gca/(2σ²gca + σ²sca)) was around unity both for Fe and Zn densities, implying preponderance of additive gene action. Further, highly significant positive correlation between mid-parent values and hybrid performance, and no correlation between mid-parent values and mid-parent heterosis confirmed again the predominant role of additive gene action for these micronutrients. Barring a few exceptions with one parent, hybrids did not outperform the parents having high Fe and Zn levels. This showed that there would be little opportunity, if any, to exploit heterosis for these mineral micronutrients in pearl millet. In general, high Fe and Zn levels in both of the parental lines would be required to increase the probability of breeding high Fe and Zn hybrids.     

Genetic mapping of quantitative trait locus for the leaf morphological traits in a recombinant inbred line population by ultra-high–density maps across multi-environments of maize (Zea mays)

Leaf is the main organ of photosynthetic reaction of plants. Studying the genetic mechanism that affects the leaf shape is very important for the improvement of maize production. In this study, a RIL population, derived from a cross between Ye478 and Qi319, was planted in four different environments, and six leaf morphological traits were measured, including the leaf angle of first leaf above ear, the leaf angle of first leaf below ear, leaf orientation value, leaf area of first leaf above ear, leaf area of ear and leaf area of first leaf below ear. By combining with a genetic map containing 4,602 bin markers, 39 QTLs associated with leaf morphological traits were identified. Among them, four QTLs explained more than 10% of the phenotypic variance, and the QTL qLOV8-2 which controlled LOV not only had a phenotypic contribution rate of 13.86% but also was detected in four environments, which could be considered as a stable major QTL. These results provide useful information for understanding the molecular mechanisms controlling maize leaf morphological traits.     

Genetic analysis and genotype × environment (G × E) for grey leaf spot disease resistance in elite African maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.) germplasm

Maize grey leaf spot (GLS) disease remains an important foliar disease in sub-Saharan Africa accounting for more than 25% yield losses in maize. Information on inheritance of GLS resistance of germplasm adapted to African environments is required in new sources being identified. Therefore, hybrids generated from a 10 × 10 half-diallel mating of tropical advanced maize inbred lines were evaluated in six environments to determine combining ability, genotype × environment interaction (G × E) and the impact of GLS disease on grain yield. General combining ability effects were highly significant and accounted for 72 and 68% of the variation for GLS resistance and grain yield, respectively. Significant specific combining ability effects associated with reduced disease levels were observed in some hybrids when one parent was resistant, and these may be exploited in developing single cross maize hybrids. Regression analysis showed a 260–320 kg ha^?1 decrease in maize grain yield per each increase in GLS disease severity score, and significant associations (r = ?0.31 to ?0.60) were observed between grain yield and GLS severity scores. This showed the potential of GLS disease to reduce yield in susceptible varieties grown under favourable disease conditions, without control measures. Genotype and genotype × environment biplots and correlation analysis indicated that the significant G × E observed was not due to changes in hybrid ranking, implying absence of a significant crossover interaction. Therefore, predominance of additive gene effects imply that breeding progress for GLS disease resistance would be made through selection and this could be achieved at a few hot-spot sites, such as Baynesfield and Cedara locations in South Africa, and still deploy the resistant germplasm to other environments in which they are adapted.     

QTL examination of a bi-parental mapping population segregating for “short-stature” in hop (<Emphasis Type="Italic">Humulus lupulus</Emphasis> L.)

Increased labor costs and reduced labor pools for hop production necessitate the development of strategies that improve efficiency and automation of hop production. One solution for reducing labor inputs is the use of “short-trellis” hop varieties. Unfortunately, little information exists on the genetic control of this trait in hop, and there are no known molecular markers available for selection. This preliminary study was enacted to identify QTLs associated with expression of short-stature growth phenotype using SNPs identified within genome-assembled scaffolds. A bi-parental mapping population of 87 offspring was obtained from the cross, “Pioneer × 25/95/15”. Genotyping-by-sequencing was performed on parents and offspring. SNPs were identified using TASSEL v3.0 with either ‘Teamaker’ reference genome or ‘Shinsuwase’ genome. The genetic map derived from ‘Teamaker’ SNPs was far superior and was used for all further analysis. QTL analysis identified eight QTLs linked to short stature with five showing strong statistical association based upon three different statistical analyses. All eight QTLs were found on linkage group one. Evaluation of scaffolds containing SNP markers located at or surrounding QTL regions (±1 cM) identified 67 putative genes—several of which are known structural genes. A genome-wide scan of SNP markers identified an additional marker found on a scaffold containing a putative gene (Aspartyl protease family protein) known to induce dwarf characteristics in other species. Further validation of significantly associated markers on different populations is necessary prior to implementation in marker-assisted selection.