Transgressive segregation for phenological traits in barley explained by two major QTL alleles with additivity

Quantitative trait loci (QTL) analysis can contribute to a deeper understanding of crop phenology. The parents of a barley mapping population have similar growth and development profiles, but the progeny show transgressive segregation for phenological traits. These phenotypes were measured in eight field experiments, using different planting dates over 3 years. Five QTL, on four chromosomes, were detected for anthesis date. The four maturity QTL were on the same chromosomes as the anthesis QTL. Five QTL for grain filling were detected on all chromosomes. Three QTL, on chromosomes 1H and 2H, were detected for photoperiod sensitivity. Both parents contributed higher value alleles for all traits, except for photoperiod sensitivity. QTL epistasis was not significant. Two QTL explained most of the phenotypic variation for anthesis and physiological maturity. Non‐parental combinations of alleles at these loci account for the phenotypic transgressive segregation. Candidate genes for these QTL effects are eps2S (2H) and denso (3H). QTL for other traits had smaller effects and were coincident with genes and/or QTL for the same traits reported in other germplasm.     

Quantitative trait loci for fruit size and flowering time-related traits under domestication and diversifying selection in cucumber (Cucumis sativus)

In cucumber, the genetic basis of traits under domestication and/or diversifying selection is not well understood. Here, we reported QTL mapping for flowering time and fruit size-related traits with segregating populations derived from a cultivated × wild cross. Phenotypic data of flowering time (FT), fruit size (FS), fruit number (FN) and fruit weight per plant (FW) were collected in multiple environments. QTL analysis identified 19 QTL for these traits. We found that the major-effect QTL FT1.1 played an important role in regulating flowering time in cultivated cucumber, whereas the minor-effect QTL FT6.3 contributed to photoperiod sensitive flowering time during domestication. Two novel consensus FS QTL, FS1.4 and FS2.3, seem to be the targets of selection during breeding for the US processing cucumber. All other FS QTL were co-localized with previously detected QTL using populations derived from cultivated cucumbers, suggesting that they were under selection during both initial domestication and subsequent improvement. Results from this study also suggested that the wild cucumber is a useful resource for capturing positive transgressive segregation and novel alleles that could be explored in cucumber breeding.     

Phenotypic and QTL analyses of herbage production-related traits in perennial ryegrass (<Emphasis Type="Italic">Lolium perenne</Emphasis> L.)

Phenotypic and genetic evaluation of morphological traits associated with herbage biomass production was undertaken in a perennial ryegrass (Lolium perenne L.) biparental F₁ mapping population (n = 200) with parent plants from cultivars ‘Grasslands Impact’ and ‘Grasslands Samson’. Morphological traits measured on three clonal replicates of the parental genotypes and 200 F₁ progeny in a glasshouse in two separate trials (autumn and spring) included: dry weight (DW), leaf elongation rate (LER), initial tiller number (TNs), final tiller number (TNe), site filling (Fs), tiller weight (TW), leaf lamina length, leaf tip and ligule appearance rates (ALf, ALg) and leaf elongation duration (LED). Principal component analysis of patterns of trait association identified negative correlation between TNs or TNe, and TW as the primary basis for morphological difference and indicated that either high LER or long LED could reduce TN. Plants with higher LER tended to have increased DW. Quantitative trait loci (QTL) were detected on all seven linkage groups (LG) of a perennial ryegrass linkage map for all but three traits. A total of 61 QTL were identified, many of which clustered at 15 shared genome locations. Significant genotype by environment effects were encountered, evidenced both by variation between experiments in genotype rankings and by a general lack of commonality for QTL for the same traits in the different experiments. Only five QTL, for ALf, ALg and TN, were conserved between autumn and spring trials. A QTL for TN and DW on LG6 is a strong candidate for application of MAS in future plant improvement work and was found to be co-linear with QTL for equivalent traits reported on chromosome 2 in rice.     

Dent corn genetic background influences QTL detection for grain yield and yield components in high-oil maize

Despite the well-recognized importance of grain yield in high-oil maize (Zea mays L.) breeding and production, few studies have reported the application of QTL mapping of such traits. An inbred line of high-oil maize designated ‘GY220’ was crossed with two dent maize inbred lines to generate two connected F_2:3 populations with 284 and 265 F_2:3 families. Our main objective was to evaluate the influence of genetic background on QTL detection of grain yield traits through comparisons between the F_2:3 populations. The field experiments were conducted during the spring in Luoyang and summer in Xuchang, Henan, China. Two genetic linkage maps were constructed with a genetic distance of 2111.7 and 2298.5 cM using 185 and 173 polymorphic SSR markers, respectively. In total, 18 and 15 QTL were detected for six grain yield traits in the two populations. Only one common QTL marker was shared between the two populations. A QTL cluster associated with five traits was identified at bin 1.05–1.06, including the shared QTL for 100GW, which demonstrated the largest effect (16.7%). Among the detected QTL, 12 digenic interactions were identified. Our results reflect the substantial influence of dent maize genetic background on QTL detection of grain yield traits.     

QTL mapping of improving forage maize starch degradability in European elite maize germplasm

Improving maize starch content is of great importance for both forage and grain yield. In this study, 13 starch degradability traits were analysed including percentage of the seedling area, floury endosperm, hard endosperm of total grain area, percentage of the floury endosperm surface and vitreousness ratio surface hard: floury endosperm surface, etc. We mapped quantitative trait loci (QTL) in a biparental population of 309 doubled haploid lines based on field phenotyping at two locations. A genetic linkage map was constructed using 168 SSR (simple sequence repeat) markers, which covered 1508 cM of the maize genome, with an average distance of 9.0 cM. Close phenotypic and genotypic correlations were found for all traits, and were all statistically significant (p = 0.01) at two locations. Major QTL for more than two traits were detected, especially in two regions in bins 4.05–4.06 and 7.04–7.05, associated with 13 and 9 traits, respectively. This study contributes to marker‐assisted breeding and also to fine mapping candidate genes associated with maize starch degradability.     

QTL mapping of a partial resistance to the corn delphacid‐transmitted viruses in Lepidopteran‐resistant maize line Mp705

A partial resistance to maize mosaic virus (MMV) and maize stripe virus (MStV) was mapped in a RILs population derived from a cross between lines MP705 (resistant) and B73 (susceptible). A genetic map constructed from 131 SSR markers spanned 1399 cM with an average distance of 9.6 cM. A total of 10 QTL were detected for resistance to MMV and MStV, using composite interval mapping. A major QTL explaining 34–41% of the phenotypic variance for early resistance to MMV was detected on chromosome 1. Another major QTL explaining up to 30% of the phenotypic variation for all traits of resistance to MStV was detected in the centromeric region of chromosome 3 (3.05 bin). After adding supplementary SSR markers, this region was found to correspond well to the one where a QTL of resistance to MStV already was located in a previous mapping study using an F₂ population derived from a cross between Rev81 and B73. These results suggested that these QTL of resistance to MStV detected on chromosome 3 could be allelic in maize genome.     

Mapping QTLs of traits contributing to yield and analysis of genetic effects in tetraploid cotton

Fiber yield and yield components – including lint index (LI), seed index (SI), lint yield (LY), seed cotton yield (SCY) and number of seeds per boll (NSPB) – were investigated on the farm of Huazhong Agricultural University in a population of 69 F₂ individuals and corresponding F_2:3 families derived from a cross between high-fiber-yield Gossypium hirsutum CV Handan 208 and a low-fiber-yield Gossypium barbadense CV Pima 90. On the basis of the genetic map constructed previously from the same population by Lin et al. (Plant Breed., 2005), quantitative trait locus (QTL) analysis was performed with the software QTL Cartographer V2.0 using composite interval mapping method (LOD ≥ 3.0). A total of 21 QTLs were identified, which were located in 15 linkage groups. The number of QTLs per trait ranged from one to seven. Of these QTLs detected, one affecting LI explained 24.3% of phenotypic variation (PV), five influencing SI explained 16.15–39.21% of PV, seven controlling LY explained 13.01–28.35% of PV, and two controlling SCY explained 22.76 and 39.97% of PV, respectively. Simultaneously, the detected six QTLs for NSPB were located on five linkage groups, which individually explained 28.01–38.32% of the total phenotypic variation. The results would give breeders further insight into the genetic basis of fiber yield.     

Efficiency of low heritability QTL mapping under high SNP density

The efficiency of quantitative trait locus (QTL) mapping methods needs to be investigated assuming high single nucleotide polymorphism (SNP) density and low heritability QTLs. This study assessed the efficiency of the least squares, maximum likelihood, and Bayesian approaches for QTL mapping assuming high SNP density and low heritability QTLs. We simulated 50 samples of 400 F₂ individuals, which were genotyped for 1000 SNPs (average density of one SNP/centiMorgan) and phenotyped for three traits controlled by 12 QTLs and 88 minor genes. The genes were randomly distributed in the regions covered by the SNPs along ten chromosomes. The QTL heritabilities ranged from approximately 1–2% and the sample sizes were 200 and 400. The power of QTL detection ranged from 30 to 60%, the false discovery rate (FDR) ranged from only 0.5–1.2%, and the bias in the QTL position ranged from 4 to 6 cM. The QTL mapping efficiency was not influenced by the degree of dominance. The statistical approaches were comparable regarding the FDR. Regression-based and simple interval mapping methods showed equivalent power of QTL detection and mapping precision. Compared to interval mapping, the inclusive composite interval mapping provided slightly greater QTL detection power and mapping precision only for the intermediate and high heritability QTLs. By maximizing the prior number of QTLs, the Bayesian analysis provided the greatest power of QTL detection. No method proved to be superior.     

Quantitative trait loci analysis of stem strength and related traits in soybean

Stem strength is one of the major influencing factors of lodging in soybean [Glycine max (L.) Merr.] as well as other crops. To identify quantitative trait loci (QTL) associated with stem strength and related traits in soybean, a recombinant inbred line (RIL) population consisting of 165 lines derived from Zhongdou No. 29 × Zhongdou No. 32 was used in 3 years. Significant positive correlations were found among the four traits (stem strength, stem diameter, number of nodes, root dry weight). A linkage map spanning 1,240.7 cM was constructed using 245 SSR (simple sequence repeat) markers and a phenotypic marker (leaflet shape). By composite interval mapping and two-round strategy of QTL meta-analysis, 32 consensus QTL and 19 unique QTL were identified, respectively. Of eight pleiotropic unique QTL, two QTL (uq.A2-2 and uq.A2-3) located at the intervals of 23.2–26.8 and 38.5–42.4 cM on linkage group A2, respectively, were associated with all the four traits. Additive × environment (ae) interaction effects, epistasis (aa) and epistasis × environment (aae) interaction effects of QTL were detected as well. The results provide useful information for further genetic studies on stem strength of soybean.     

QTL mapping of yield-related traits in the wheat germplasm 3228

The new wheat germplasm 3228, a putative derivative of tetraploid Agropyron cristatum Z559 and the common wheat Fukuhokomugi, has superior features in yield-related traits, particularly in spike morphological traits, such as large spike and superior grain number. To identify favorable alleles of these traits in 3228, 237 F_2:3 families were developed from the cross 3228/Jing 4839. A genetic map was constructed using 179 polymorphic SSR and EST-SSR markers. A total of 76 QTL controlling spike number per plant (SNP), spike length (SL), spikelet number per spike (SNS), floret number per spikelet (FNS), grain number per spike (GNS) and thousand-grain weight (TGW) were detected on 16 chromosomes. Each QTL explained 1.24–27.01% of the phenotypic variation, and 9 QTL (28.95%) were detected in two or all environments. Additive effects of 45 QTL were positive with 3228 alleles increasing the QTL effects, 31 QTL had negative effects indicating positive contributions from Jing 4839. Three important clusters involving all traits were located on chromosomes 5A, 6A and 4B, and several co-located QTL were also found. Most of the QTL detected on the three chromosome regions could contribute to the use of 3228 in breeding for grain yield improvement.     

基于贝叶斯统计的谷物胚乳性状QTL多区间作图方法

贝叶斯统计学已被广泛地应用在现代科学的各个研究领域。本研究将贝叶斯统计方法和谷物三倍体胚乳性状数量遗传模型相结合,以F₂群体中各植株的分子标记基因型以及植株上若干粒自交种子胚乳性状的单粒观测值为数据模式,提出了胚乳数量性状基因座(QTL)多区间作图的贝叶斯方法。该方法首先构建胚乳性状的多区间多QTL遗传模型,然后通过基于Gibbs抽样和Metropolis-Hastings算法实现的马尔可夫链蒙特卡罗(MCMC)方法同时获得多个QTL效应和位置的估计。方法的有效性通过一条长染色体的模拟实验进行了验证,结果表明,本文提出的贝叶斯多区间方法能够准确地估计胚乳性状QTL的位置和效应,并可有效区分两种显性效应。     

Mapping quantitative trait loci associated with starch paste viscosity in rice (Oryza sativa L.) under different environmental conditions

A recombinant inbred line (RIL) population consisting of 153 lines derived from a cross between indica ‘Zhongyouzao8’ and japonica ‘Toyonishiki’ rice was used to detect stable quantitative trait loci (QTL) for rapid visco analyser (RVA) characteristics under four environmental conditions. We identified 93 QTL for RVA profiles, and four pleiotropic regions harbouring stably expressed QTL were detected on chromosomes 2, 6, 7 and 11. These newly identified and stable QTL will facilitate further research into the genetic mechanism regulating RVA profiles. Amylose content (AC) was correlated with RVA traits. AC and RVA traits were unaffected by indica–japonica subspecies differentiation, suggesting that RVA profiles were mainly influenced by the Wx gene. The RIL population was divided into two subpopulations according to Wx genotypes. A total of 106 QTL associated with RVA profiles were detected in the subpopulations. These QTL differed from those detected in the whole population in terms of their genomic location, number, logarithm of odds values and amount of phenotypic variance explained. Using this strategy, we detected QTL with minor effects and eliminated false due to the Wx gene.     

QTL mapping of nutrient components in maize kernels under low nitrogen conditions

A set of 213 F_2:3 families were used to investigate the effects of nitrogen (N) on grain yield and the concentrations of three nutrient components in maize (Zea mays L.) kernels. A genetic linkage map was constructed using 189 SSR (simple sequence repeat) markers, spanning a total of 2003 cM, including 11 linkages, and the families were evaluated under high N and low N conditions at two farm locations. The results indicate that low N conditions may induce an increase in starch concentration, but a decrease in protein levels. Twenty‐six quantitative trait loci (QTL) were detected for four measured traits in the two N treatments at both locations, including eight QTL for grain yield, seven QTL for oil content, six QTL for protein content and five QTL for starch content. The total number of QTL detected for the four measured traits under high N levels was greater than the QTL detected under low N conditions, and several QTL were identified that specifically expressed under different N conditions. These particular QTL could help provide greater understanding of the genetic basis of N‐usage efficiency.     

Genetics and mapping of quantitative traits for nodule number, weight, and size in soybean (Glycine max L.[Merr.])

Soybean research has found that nodule traits, especially nodule biomass, are associated with N₂ fixation ability. Two genotypes, differing in nodule number per plant and individual nodule weight, KS4895 and Jackson, were mated to create 17 F₃- and 80 F₅-derived RILs. The population was mapped with 664 informative markers with an average distance of less than 20 cM between adjacent markers. Nodule traits were evaluated in 3-year field trials. Broad-sense heritability for nodule number (no. plant^?1), individual nodule dry weight (mg nodule^?1), individual nodule size (mm nodule^?1), and total nodule dry weight (g plant^?1) was 0.41, 0.42, 0.45, and 0.27, respectively. Nodule number was negatively correlated with individual nodule weight and size. Nodule number, individual nodule weight, and size are major components which likely contributed to increased total nodule weight per plant. Composite interval mapping (CIM) identified eight QTLs for nodule number with R² values ranging from 0.14 to 0.20. Multiple interval mapping (MIM) identified two QTLs for nodule number, one of which was located close to the QTL identified with CIM. Six QTLs for individual nodule weight were detected with CIM, and one QTL was identified with MIM. For nodule size, CIM identified seven QTLs with R² values ranging from 0.14 to 0.27. Five QTLs for total nodule weight were detected with CIM, one of which was located close to a QTL identified with MIM. These results document the first QTL information on nodule traits in soybean from field experiments utilizing a dense, complete linkage map.     

Dynamic QTL mapping for plant height in the hybrid population of Agropyron Gaertn.

Plant height (PH) plays a very important role for plant breeding and also serves as a model trait to dynamic development study. The dynamic quantitative trait locus (QTL) analysis for PH of Agropyron Gaertn. was carried out in a cross-pollination (CP) hybrid population of A. Gaertn. based on the phenotypic data of PH at different developmental stages in four year-sites and the constructed single-nucleotide polymorphism (SNP) genetic map. The results showed that 69 QTL and nine major QTL were detected by unconditional QTL mapping. A total of 107 QTL and 13 major QTL were found using conditional QTL mapping. Forty-seven QTL were detected by two methods. Two unconditional QTL, Qph2-5 and Qph3-4, were expressed as major and stable QTL for PH. Four major stable conditional QTL for PH, cQph4-1, cQph4-8, cQph6-2 and cQph6-7, were detected. Two conditional PH QTL, cQph3-5 and cQPh3-7, were identified in four environments over multiple stages. The gene/QTL controlling PH was expressed in a certain spatiotemporal manner. These results could also provide a reference for genetic and breeding research of related plant.     

Molecular mapping of quantitative trait loci for plant growth, yield and yield related traits across three diverse locations in a doubled haploid rice population

A doubled haploid (DH) population of 125lines derived from IR64 × Azucena, an indica – japonica cross were grown in three different locations in India during the wet season of 1995. The parents of mapping population had diverse phenotypic values for the eleven traits observed. The DH lines exhibited considerable amount of variation for all the traits. Transgressive segregants were observed. Interval analysis with threshold LOD > 3.00 detected a total of thirty four quantitative trait loci (QTL) for eleven traits across three locations. The maximum number of twenty QTL were detected at Punjab location of North India. A total of seven QTL were identified for panicle length followed by six QTL for plant height. Eight QTL were identified on three chromosomes which were common across locations. A maximum of seven QTL were identified for panicle length with the peak LOD score of 6.01 and variance of 26.80%. The major QTL for plant height was located on Chromosome 1 with peak LOD score of 16.06 flanked by RZ730-RZ801 markers. Plant height had the maximum number of common QTL across environment at the same marker interval. One QTL was identified for grain yield per plant and four QTL for thousand grain weight. Clustering of QTL for different traits at the same marker intervals was observed for plant height, panicle exsertion, panicle number, panicle length and biomass production. This suggests that pleiotropism and or tight linkage of different traits could be the plausible reason for the congruence of several QTL. Common QTL identified across locations and environment provide an excellent opportunity for selecting stable chromosomal regions contributing to yield and yield components to develop QTL introgressed lines that can be deployed in rice breeding program. This revised version was published online in July 2006 with corrections to the Cover Date.     

Detecting major QTL associated with resistance to bacterial blight using a set of rice reciprocal introgression lines with high density SNP markers

Bacterial blight (BB), caused by Xanthomonas oryzae pv. oryzae (Xoo), is a worldwide rice disease. QTL providing BB resistance were identified using a set of introgression lines (ILs) derived from a cross between Teqing and Lemont, and 3924 evenly distributed SNP markers developed from the two parents. After inoculating three Xoo pathotypes, CII, CIV and CV, seven major BB resistance‐associated QTL were detected. The alleles at all loci with improved BB resistance were from the Teqing background. Four QTL resistance to CII and CIV were identified in the reciprocal backgrounds across 2 years. Six QTL conferring resistance to CV identified in Teqing background were not detected in Lemont background, indicating that genetic background had a strong effect on the BB resistance‐associated QTL. Based on the interactions of 27 significant digenic QTL pairs among the seven main‐effect QTL, the QTL were divided into three strain‐specific groups. Genotype analyses of resistant ILs suggest that rice lines with a high level of resistance to BB can be achieved by pyramiding R gene(s) and QTL that interact with R gene(s).     

Quantitative trait loci mapping of forage stover quality traits in six mapping populations derived from European elite maize germplasm

Four forage maize stover quality traits were analysed including in vitro digestibility of organic matter (IVDOM), neutral detergent fibre (NDF), water‐soluble carbohydrates (WSC) and digestibility of NDF (DNDF). We mapped quantitative trait loci (QTL) in three DH (doubled haploid) populations (totally 250–720 DH lines): one RIL population (358 lines) and two testcross (TC) populations, based on field phenotyping at multiple locations and years for each. High phenotypic and genotypic correlations were found for all traits and significant (p < .01) at two locations, and NDF was negatively correlated with the other traits. QTL analyses were conducted by composite interval mapping. A total of 33, 23, 32 and 25 QTL were identified for IVDOM, NDF, WSC and DNDF, respectively, with three, four, five and two major QTL for each. Few consistent QTL for IVDOM, WSC and DNDF were detected in more than two populations. This study contributed to the identification of key QTL associated with forage maize digestibility traits and is beneficial for marker‐assisted breeding and fine mapping of candidate genes associated with forage maize quality.     

QTL detection for grain oil and starch content and their associations in two connected F<Subscript>2:3</Subscript> populations in high-oil maize

High-oil maize (Zea mays L.) has special value in animal feed and human food. Two hundred and eight-four and 265 F_2:3 families developed from two crosses between one high-oil maize inbred and two normal dent maize inbreds were evaluated for grain oil and starch contents under two environments. Using composite interval mapping, 1–6 QTL for each trait were detected under each environment and in combined analysis in both populations. Only one common QTL across two environments in each population and across two populations were found for starch content. Among the detected QTL, nine digenic interactions with small effects were identified. Comparison of single-trait QTL and the results of multiple-trait QTL mapping showed that oil content might be complicatedly correlated with starch content. Although single-trait QTL with the same parental effects for both traits and oil-starch QTL were all detected at the same genetic bin 6.04 as the cloned high-oil QTL (qHO6) with no unfavorable effects on grain weight, our results did reflect the difficulty to realize simultaneous improvement on grain oil and starch contents. Of course, these results should be validated in further experiments under more environments using RILs, NILs and other permanent populations.