Quantitative trait loci for agronomic traits in soybean

There continues to be improvement in seed yields of soybean by conventional breeding, but molecular techniques may provide faster genetic gains. The objective of this study was to identify quantitative trait loci (QTL) associated with the agronomic traits seed yield, lodging, plant height, seed filling period and plant maturity in soybean. To achieve this objective, 101 F₆‐derived recombinant inbred lines (RIL) from a population developed from a cross of N87‐984‐16 × TN93‐99 were used. Experiments were conducted in six environments during 2002–2003. Heritability estimates on an entry mean basis from data combined across environments ranged from 0.12 to 0.65 for seed yield and seed filling period, respectively. Composite interval mapping detected one QTL for yield (near Satt076), two for lodging (near Satt225 and Satt593) and four for maturity (near Satt263, Satt292, Satt293 and Satt591) in this population. Additional environmentally sensitive QTL for these traits, and for seed filling period and plant height are also reported. The QTL associated with agronomic traits that we report and the recently released germplasm (PI 636460) from this population may be useful in soybean breeding programmes.     

Quantitative trait loci mapping for yield-related traits under low and high planting densities in maize (Zea mays)

Average maize yield per hectare has increased significantly because of the improvement in high-density tolerance, but little attention has been paid to the genetic mechanism of grain yield response to high planting density. Here, we used a population of 301 recombinant inbred lines (RILs) derived from the cross YE478 × 08–641 to detect quantitative trait loci (QTLs) for 16 yield-related traits under two planting densities (57,000 and 114,000 plants per ha) across four environments. These yield-related traits responded differently to high-density stress. A total of 110 QTLs were observed for these traits: 33 QTLs only under low planting density, 50 QTLs under high planting density and 27 QTLs across both densities. Only two major QTLs, qCD6 and qWKEL2-2, were identified across low- and high-density treatments. Seven environmentally stable QTLs were also observed containing qED6, qWKEL3, qRN3-3, qRN7-2, qRN9-2 and qRN10 across both densities, as well as qRN9-1 under low density. In addition, 16 and eight pairs of loci with epistasis interaction (EPI) were detected under low and high planting densities, respectively. Additionally, nine and 17 loci showed QTL × environment interaction (QEI) under low- and high-density conditions, respectively. These interactions are of lesser importance than the main QTL effects. We also observed 26 pleiotropic QTL clusters, and the hotspot region 3.08 concentrated nine QTLs, suggesting its great importance for maize yield. These findings suggested that multiple minor QTLs, loci with EPI and QEI, pleiotropy and the complex network of “crosstalk” among them for yield-related traits were greatly influenced by plant density, which increases our understanding of the genetic mechanism of yield-related traits for high-density tolerance.     

Inheritance of firmness in raw cucumber (Cucumis sativus L.) fruit

Summary Inheritance of raw cucumber fruit texture (Magness-Taylor Fruit Pressure Tester firmness) was investigated over a 4-year period from 1971–1974. Results from 2 separate but related experiments suggested that firmness was quantitatively inherited with sufficient additive effects to permit gain from selection. In a selection study within 4 F₂ populations derived from crosses between firm (Chipper and Gy3) and soft (Mincu and Green F) fruit type cultivars, variation among and within F₃ and F₄ families was significant but overall family means were not significantly higher than the high parent in any of the 4 crosses. Narrow sense heritability estimates for fruit texture were 0.80 in the Mincu × Chipper population and 0.77 in the Green F × Chipper, Mincu × Gy3, and Gy3 × Green F crosses. In a separate experiment, generation means analysis was used to assess the mode of gene action in 2 crosses: Green F × Chipper, and Gy3 × Green F. Additive genetic effects accounted for 98.8% and 99.3% of the total genetic variation within each cross, respectively.Scientific Journal Series Paper No. 9794.     

Mapping quantitative trait loci for yield-related traits in soybean (Glycine max L.)

Development of soybean cultivars with high seed yield is a major focus in soybean breeding programs. This study was conducted to identify genetic loci associated with seed yield-related traits in soybean and also to clarify consistency of the detected QTLs with QTLs found by previous researchers. A population of 135 F_2:3 lines was developed from a cross between a vegetable soybean line (MJ0004-6) and a landrace cultivar from Myanmar ({"type":"entrez-nucleotide","attrs":{"text":"R18500","term_id":"772110","term_text":"R18500"}}R18500). They were evaluated in the experimental field of Kasetsart University, Kamphaeng Saen, Nakhon Pathom, Thailand in a randomized complete block design with two replications each in 2011 and 2012 growing seasons. The two parents exhibited contrasting characteristics for most of the traits that were mapped. Analysis of variance showed that the main effects of genotype and environment (year) were significant for all studied traits. Genotype by environment interaction was also highly significant for all the traits. The population was genotyped by 149 polymorphic SSR markers and the genetic map consisted of 129 SSR loci which converged into 38 linkage groups covering 1156 cM of soybean genome. There were 10 QTLs significantly associated with seed yield-related traits across two seasons with single QTLs explaining between 5.0% to 21.9% of the phenotypic variation. Three of these QTLs were detected in both years for days to flowering, days to maturity and 100 seed weight. Most of the detected QTLs in our research were consistent with earlier QTLs reported by previous researchers. However, four novel QTLs including SF1, SF2 and SF3 on linkage groups L and N for seed filling period and PN1 on linkage group D1b for pod number were identified in the present study.     

Novel quantitative trait loci for yield and yield related traits identified in Basmati rice (Oryza sativa)

Increasing crop productivity is one of the prime goals of crop breeding research. Rice grain yield is a complex quantitative trait governed by polygenes. Although several QTLs governing grain yield traits have been reported and limited attempts have been made to map QTLs for grain yield parameters in Basmati rice. A population from the cross Sonasal and Pusa Basmati 1121 comprising 352 RILs was generated through the single seed descent method. A total of 12 QTLs governing yield and yield-related traits were mapped on six chromosomes, namely, 1, 2, 3, 7, 8 and 9, of which five QTLs were novel. We identified a novel and robust epistatic QTL (qPH1.1 and qPL1.1) governing plant height and panicle length, flanked by the markers RM5336-RM1 on chromosome 1. The gene encoding brassinosteroid insensitive 1-associated receptor kinase 1 precursor is the putative candidate gene underlying this epistatic QTL. Another novel QTL, qNT3.1, governing tiller number was bracketed to a region of .77 Mb between the markers RM15247 and RM15281 on chromosome 3. Of the 57 annotated gene models, Os03g0437600 encoding alpha/beta-fold hydrolase, a homologous to AtKai2 is a putative candidate gene underlying the novel QTL qNT3.1. The other QTLs such as qDFF1.1 governing days to 50% flowering co-localizes with the gene Ghd7, QTL for plant height qPH1.2 co-localizes with the gene sd1, the QTLs for panicle length co-localizes with FUWA and DEP2, the QTL for tiller number co-localizes with OsRLCK57 and QTLs for thousand-grain weight co-localize with the major gene GS3. The QTLs identified in the current study can be effectively used in marker-assisted selection for developing Basmati rice varieties with a higher yield.     

Sample size required for marker assisted selection in improving quantitative traits of self-fertilizing species

The efficiency of selection for desired trait genotypes in a molecular marker assisted selection for a quantitative trait in self-fertilizing crop is considered. The QTLs controlling the trait were assumed to be unlinked. It was supposed that the selection starts in F₂, derived from a cross between inbred lines, and this selection will terminate if one or more plants with the desired trait genotype is found. If no plant with the desired trait genotype is found in F₂ then the selection is continued in the F₃ progeny that is derived from a single selected F₂ plant. Which F₂ plant is to be selected was determined according to the rank which is related to the marker genotype of the F₂ plants. And this rank was based on the expected frequency of the desired trait genotype in the progeny. The plant with the top rank among all F₂ plants is then selected with the first priority. Additionally the number of F₃ plants in the progeny was set to be equal to the number of plants that are required for detecting one or more plants with the desired trait genotype with a given probability. The probability of getting at least one plant with the desired trait genotype is expressed as a function of the number of F₂ plants (N).The required value for N and the total number of plants (T) in F₂ and F₃ for detecting at least one plant with the desired trait genotype were calculated for different situations. T was always smaller for a single marker than for flanking markers. The minimum of T and monotonous decrease of N can be observed when the cumulative-expected-frequency of selected marker genotypes of F₂ plants increased. This revised version was published online in July 2006 with corrections to the Cover Date.     

Marker-assisted selection for complex trait in common bean (Phaseolus vulgaris L.) using QTL-based index

A procedure was developed for marker-assisted selection of complex traits for common bean (Phaseolus vulgarisL.) using an index based on QTL-linked markers and ultrametric genetic distances between lines and a target parent. A comparison of the mean seed yields of the top five lines selected by different schemes demonstrated that the highest yielding group was selected on the basis of a combination of phenotypic performance and a high QTL-based index,followed by groups identified by a high QTL-based-index, conventional selection,and a low QTL-based-index. This study demonstrated a simple way to use information obtained from QTL studies to make selection decisions. The study also showed that the use of the QTL-based-index in conjunction with the ultrametric genetic distance to the target parent would enablea plant breeder to select lines that retain important QTL in a desirable genetic background. Therefore, this type of MAS would be expected to be superior to the phenotypic selection. This revised version was published online in August 2006 with corrections to the Cover Date.     

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

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

An introduction to markers, quantitative trait loci (QTL) mapping and marker-assisted selection for crop improvement: The basic concepts

Recognizing the enormous potential of DNA markers in plant breeding, many agricultural research centers and plant breeding institutes have adopted the capacity for marker development and marker-assisted selection (MAS). However, due to rapid developments in marker technology, statistical methodology for identifying quantitative trait loci (QTLs) and the jargon used by molecular biologists, the utility of DNA markers in plant breeding may not be clearly understood by non-molecular biologists. This review provides an introduction to DNA markers and the concept of polymorphism, linkage analysis and map construction, the principles of QTL analysis and how markers may be applied in breeding programs using MAS. This review has been specifically written for readers who have only a basic knowledge of molecular biology and/or plant genetics. Its format is therefore ideal for conventional plant breeders, physiologists, pathologists, other plant scientists and students.     

Quantitative trait loci for leafhopper (Empoasca fabae and Empoasca kraemeri) resistance and seed weight in the common bean

A population of 108 common bean recombinant inbred lines (RILs) (F_5:6‐9), derived from a leafhopper (Empoasca fabae and E. kraemeri)‐susceptible cultivar (‘Berna’) and a leafhopper‐resistant line (EMP 419) was used to identify molecular markers genetically linked to leafhopper resistance and seed weight. Bulked segregant analysis and quantitative trait analysis identified eight markers that were associated with resistance to E. fabae, and four markers that were associated with E. kraemeri resistance. Three markers were associated with resistance to both species. A partial linkage map of the bean genome was constructed. Composite interval mapping identified quantitative trait loci (QTL) for resistance to both leaf hopper species on core‐map linkage groups B1, B3 and B7. QTL for seed weight were found close to the locus controlling testa colour and an α‐phaseolin gene.     

Quantitative trait loci (QTL) mapping of silique length and petal colour in Brassica rapa

Recombinant inbred lines (RILs) derived from a cross between Brassica rapa L. cv. ‘Sampad’, and an inbred line 3‐0026.027 was used to map the loci controlling silique length and petal colour. The RILs were evaluated under four environments. Variation for silique length in the RILs ranged from normal, such as ‘Sampad’, to short silique, such as 3‐0026.027. Three QTL, SLA3, SLA5 and SLA7, were detected on the linkage groups A3, A5 and A7, respectively. These QTL explained 36.0 to 42.3% total phenotypic variance in the individual environments and collectively 32.5% phenotypic variance. No additive × additive epistatic interaction was detected between the three QTL. Moreover, no QTL × environment interaction was detected in any of the four environments. The number of loci for silique length detected based on QTL mapping agrees well with the results from segregation analysis of the RILs. In case of petal colour, a single locus governing this trait was detected on the linkage group A2.     

Quantitative trait loci for quality and agronomic traits in two advanced backcross populations in oat (Avena sativa L.)

Using the advanced backcross quantitative trait loci (AB‐QTL) strategy, we successfully transferred and mapped valuable allelic variants from the high β‐glucan (BG) accession IAH611 (PI 502955), into the genome of cultivar ‘Iltis’. By backcrossing one BC₁F₁ plant to ‘Iltis’, we developed two BC₂F_2‐6 populations A and B, comprising 98 and 72 F₂‐individuals, respectively. Genotyping of BC₂F₂ individuals with predominantly AFLP markers resulted in 12 linkage groups with a map size of 455.4 cM for Population A and 11 linkage groups with a map size of 313.5 cM for Population B. Both populations were grown at three sites in Germany over a three‐year period. Individuals were then phenotyped for 13 traits including grain yield (YD) and β‐glucan content (BG). QTL analysis via stepwise regression detected a total of 33 QTLs, most of which were clustered in three linkage groups. Two dense linkage groups A1 and B13 were found to be putatively homologous to groups KO_6 and KO_11 of the ‘Kanota’/‘Ogle’ map, respectively.     

Variation in heading date conceals quantitative trait loci for other traits of importance in breeding selection of rice

To identify quantitative trait loci (QTLs) associated with the primary target traits for selection in practical rice breeding programs, backcross inbred lines (BILs) derived from crosses between temperate japonica rice cultivars Nipponbare and Koshihikari were evaluated for 50 agronomic traits at six experimental fields located throughout Japan. Thirty-three of the 50 traits were significantly correlated with heading date. Using a linkage map including 647 single-nucleotide polymorphisms (SNPs), a total of 122 QTLs for 38 traits were mapped on all rice chromosomes except chromosomes 5 and 9. Fifty-eight of the 122 QTLs were detected near the heading date QTLs Hd16 and Hd17 and the remaining 64 QTLs were found in other chromosome regions. QTL analysis of 51 BILs having homozygous for the Koshihikari chromosome segments around Hd16 and Hd17 allowed us to detect 40 QTLs associated with 27 traits; 23 of these QTLs had not been detected in the original analysis. Among the 97 QTLs for the 30 traits measured in multiple environments, the genotype-by-environment interaction was significant for 44 QTLs and not significant for 53 QTLs. These results led us to propose a new selection strategy to improve agronomic performance in temperate japonica rice cultivars.     

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

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

Quantitative trait loci for early maturity and their potential in breeding for earliness in Brassica juncea

A doubled haploid population of Brassica juncea, developed from a cross between two parental lines differing for days to maturity, was used to study the efficiency of indirect selection for a primary trait through selection of secondary trait(s) over direct selection for the primary trait when quantitative trait loci information is available for both primary and secondary traits, and applied. Days to maturity was considered as primary trait, while days to first flowering, days to end of flowering, flowering period and plant height were considered as secondary traits. An RFLP linkage map was employed for QTL analysis of maturity and maturity-determinant traits, and a stable QTL B6 simultaneously affecting these two types of traits was identified. This linked QTL explained 11.7% phenotypic variation for days to maturity, 20.7% variation for days to first flowering, 24.3% variation for days to end of flowering and 14.4% variation for plant height. Phenotypic evaluation of maturity and/or maturity-determinant traits, viz. days to first flowering, days to end of flowering and plant height revealed that limited genetic advance for early maturity can be achieved through phenotypic selection of the primary and/or the secondary trait(s). However, the estimates of genetic advance for early maturity based on combined phenotypic evaluation and linked QTL data was found to be, at least, three times higher compared to genetic advance based on phenotypic evaluation only, demonstrating the potential of marker-assisted selection in breeding for early maturity in B. juncea.     

Occurrence and chromosome number of Cucumis sativus var. hardwickii (Royle) Alef. in South India and its bearing on the origin of cultivated cucumber

Summary Cucumis sativus var. hardwickii (Royle) Alef., the wild progenitor of cultivated C. sativus is reported for the first time from peninsular India. The South Indian specimens showed n=7 bivalents in PMCs. The discontinous occurrence of the wild taxon in the Himalayan regions and peninsular hills and the existence of cultivars of C. sativus adapted to the tropical and temperate climates suggest polytopic domestication of the cultivated forms. The possibility of utilizing this wild germplasm for crop improvement is indicated.     

Quantitative trait loci (QTL) affecting sugars, organic acids and other biochemical properties possibly contributing to flavor, identified in four advanced backcross populations of tomato

Although the Advanced Backcross strategy has proven very useful for QTL detection in tomato, it has been used mainly in identifying QTL for agronomic traits such as yield, color, etc. Tomato flavor is an important quality characteristic, yet it has been difficult to assess flavor or traits that affect it. In this study the AB-QTL strategy was applied to four advanced backcross populations to identify QTL for biochemical properties that may contribute to the flavor of processed tomatoes, such as sugars and organic acids. A total of 222 QTL were identified for 15 traits, including flavor as assessed by a taste panel. Correlations of certain biochemicals with flavor and possible methods of assessing and improving flavor are discussed. In particular, QTL with very significant effects associated with the ratio of sugars/glutamic acid, a trait highly correlated with improved flavor, have been identified as good targets for future work in improving the flavor of tomatoes. This revised version was published online in August 2006 with corrections to the Cover Date.     

Mapping quantitative trait loci for salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.)

Quantitative trait loci (QTLs) controlling salt tolerance at germination and the seedling stage in barley (Hordeum vulgare L.) were identified by interval mapping analysis using marker information from two doubled haploid (DH) populations derived from the crosses, Steptoe/Morex and Harrington/TR306. Interval mapping analysis revealed that the QTLs for salt tolerance at germination in the DH lines of Steptoe/Morex were located on chromosomes 4 (4H), 6(6H), and 7(5H), and in the DH lines of Harrington/TR306 on chromosomes 5(1H) and 7(5H). In both DH populations, the most effective QTLs were found at different loci on chromosome 7(5H). Genetic linkage between salt tolerance at germination and abscisic acid (ABA) response was found from QTL mapping. The QTLs for the most effective ABA response at germination were located very close to those for salt tolerance on chromosome 7 (5H) in both crosses. The QTLs for salt tolerance at the seedling stage were located on chromosomes 2(2H), 5(1H), 6(6H), and 7(5H) in the DH lines of Steptoe/Morex, and on chromosome 7(5H) in the DH lines of Harrington/TR 306. Their positions were different from those of QTLs controlling salt tolerance at germination, indicating that salt tolerance at germination and at the seedling stage were controlled by different loci. This revised version was published online in July 2006 with corrections to the Cover Date.